by Mahealani Y. Kaneshiro-Pineiro Dissertation Advisor ...thescholarship.ecu.edu/bitstream/handle/10342/1745/...Dissertation Advisor: David G. Kimmel, PhD Major Department: Coastal

JELLYFISH-HUMAN INTERACTIONS IN NORTH CAROLINA

by

Mahealani Y. Kaneshiro-Pineiro

Dissertation Advisor: David G. Kimmel, PhD

Major Department: Coastal Resources Management

ABSTRACT OF DISSERTATION

This dissertation investigated potential drivers of jellyfish-human interactions in

North Carolina. Jellyfish populations and human use of coasts are increasing; therefore,

jellyfish-human interactions are poised to become more frequent. This research

investigated how abiotic variables (i.e. temperature and salinity) and wind-driven

circulation in the Neuse River Estuary influenced the distribution and abundance of the

sea nettle, Chrysaora quinquecirrha, at six recreational sites. Life history traits were also

investigated to determine if jellyfish aggregations at the recreation sites could be linked

to sexual reproduction. Finally, the human perspective on jellyfish was investigated.

One hundred eighteen people were surveyed at 25 coastal locations prone to jellyfish

occurrences. This survey used cultural consensus theory to gather perspectives of

jellyfish ecology and how jellyfish influence society from four cultural groups: fishers

(commercial and recreational), recreationists (surfers, swimmers, etc.), North Carolina

coastal researchers, and jellyfish researchers in the United States. Results show: 1)

southwest winds 3 to 8 meters per second that occurred 1 and 5 days prior to

observations resulted in more sea nettles observed at the Neuse River Estuary

recreation sites; 2) aggregations of sea nettles resulting from wind events could not be

definitively linked to sexual reproduction based on jellyfish gonad analysis; 3) cultural

perspectives of jellyfish ecology were different among groups; this was most obvious

when the role of jellyfish in food webs was evaluated. All groups shared similar societal

perspectives, including tolerance to specific numbers of jellyfish. Overall, this research

has identified physical, ecological and societal factors that influence jellyfish-human

interactions in North Carolina and these interactions appear to be mediated by several

different factors. Understanding these factors will allow for management of jellyfish-

human interactions. Recreational areas subjected to high sea nettle occurrences based

on local oceanographic conditions may employ barrier nets to decrease the frequency

of encounters. Further studies into the dominant mode of reproduction for sea nettles

may indicate which life history stage, polyp or medusa, might be the best target for

management to reduce jellyfish-human interactions. Finally, outreach education about

common misconceptions concerning jellyfish may remove some confusion surrounding

the role of these organisms in the environment.


A Dissertation

Presented To the Faculty of the Institute for Coastal Science and Policy

East Carolina University

In Partial Fulfillment of the Requirements for the Degree

Coastal Resources Management

Primary Concentration in Coastal and Estuarine Ecology

Secondary Concentrations in Coastal Geosciences and Social Science & Coastal Policy

by


May, 2013

© Mahealani Y. Kaneshiro-Pineiro, 2013


by


APPROVED BY: DIRECTOR OF DISSERTATION: ________________________________________________________ (David G. Kimmel, PhD) COMMITTEE MEMBER: __________________________________________________ (Thomas R. Allen, PhD) COMMITTEE MEMBER: _________________________________________________ (Mary Beth Decker, PhD) COMMITTEE MEMBER: _________________________________________________ (Trip Lamb, PhD) COMMITTEE MEMBER: _________________________________________________ (David Mallinson, PhD) COMMITTEE MEMBER: _________________________________________________ (Tracy Van Holt, PhD)

CHAIR OF THE DEPARTMENT OF COASTAL RESOURCES MANAGEMENT: ________________________________ (Hans Vogelsong,, PhD)

DEAN OF THE GRADUATE SCHOOL: ___________________________________________________ Paul J. Gemperline, PhD

ACKNOWLEDGEMENTS

I have profound gratitude to many people who have contributed to this effort.

First, I thank my adviser, Dr. David Kimmel, for providing me with the opportunity to gain

this tremendous education. His patience, assistance, and mentoring throughout my time

at East Carolina University has been invaluable to me. I also thank my dissertation

committee, Tom Allen, Mary Beth Decker, Trip Lamb, David Mallinson, and Tracy Van

Holt. I have learned a great deal from each of them, and will always appreciate their

guidance throughout my studies. I could not have completed this work without the

support of the Institute for Coastal Science and Policy’s staff and faculty; especially,

Hans Vogelsong, John Rummel, Kay Evans, and Cindy Harper. I had the pleasure of

working with many students, Meghan Barbee, David Buckley, Allyn Hollingsworth, Ben

Nelson, Barryn McLaughlin, Amanda Cornelsen, Jarrod Howe, Hampton Farmer,

Robert Melvin, Amy Shore, and Thomas Steelman. Our time “jelly-fishing” will always be

dear to me and I am in deep gratitude for your help with my field work. I thank Mike

Askew, Ross Pease, Ray Bullock, Dennis Foster, Will Flannery, Ryan Ellis, and Brian

Blanton for granting me on-site access to their facilities for my field research and data

acquisition from the National Weather Service and RENCI NC-CERA, respectively. I

also appreciate the support of my lab mates and fellow colleagues in Coastal

Resources Management. I am particularly grateful for the scholarly advice, comforting

spirits, and unwavering encouragement from Andrea Dell̍Apa, Chad Smith, Wendy

Klein, Cecilia Krahforst, and Devon Eulie. Thank you for being there for me and for

anything. Finally, I have the upmost gratitude for my husband, Eric Pineiro; my mother,

Lucy Kaneshiro; brother, Kamuela Kaneshiro; and the rest of my family. I simply could

not have done this without your love and thank you for always believing in me.

TABLE OF CONTENTS

Chapter 1: Introduction to dissertation.................................................................... 1

Jellyfish-human interactions.................................................................. 1

Literature cited....................................................................................... 9

List of figures.......................................................................................... 14

Figures................................................................................................... 15

Chapter 2: Local wind dynamics influences the distribution and

abundance of the sea nettle, Chrysaora quinquecirrha,

at six estuarine recreation sites in the Neuse River Estuary........................ 17

Abstract................................................................................................. 18

Introduction............................................................................................ 19

Materials and methods.......................................................................... 23

Results................................................................................................... 28

Discussion.............................................................................................. 30

Conclusions............................................................................................ 37

Literature cited........................................................................................ 40

List of tables and figures......................................................................... 48

Tables and figures.................................................................................. 53

Chapter 3: An assessment of the potential of sea nettle,

Chrysaora quinquecirrha, sexual reproduction

in the Neuse River Estuary........................................................................... 71

Abstract................................................................................................. 72

Introduction............................................................................................ 73


Results................................................................................................... 80

Discussion.............................................................................................. 82

Conclusions............................................................................................ 85




Chapter 4: Predators, stingers and economic influencers:

a cultural consensus analysis of public perception

and ecological knowledge of jellyfish............................................................ 111

Abstract................................................................................................. 112

Introduction............................................................................................ 114


Results................................................................................................... 125

Discussion.............................................................................................. 130

Conclusions............................................................................................ 137




Appendix 1.............................................................................................. 166

Chapter 5: Dissertation conclusions and management implications........................ 170

Jellyfish-human interactions in North Carolina....................................... 171


List of figures......................................................................... 184

Figures....................................................................................................185

Appendix A: IRB letter/approval for Jellyfish Survey................................................186

Appendix B: Jellyfish survey consent form ..............................................................187

Appendix C: Jellyfish survey ....................................................................................188

CHAPTER 1

Introduction to dissertation

2

Jellyfish-human interactions

I assigned the term “jellyfish-human interactions” to describe circumstances

where jellyfish are present in coastal or oceanic environments that experience

significant human use, resulting in frequent contact between jellyfish and humans.

Understanding that the type of jellyfish-human interaction is dependent on the jellyfish

species, and if the species is favorable or unfavorable to humans, it is likely that

jellyfish-human interactions vary globally. For example, many jellyfish species are edible

(Hsieh et al. 2001) and jellyfish fisheries worldwide harvest more than 500,000 tons of

jellyfish annually (Pitt 2010). In Palau, tourists will travel to “Jellyfish Lake” to swim with

large numbers of Mastigias sp. jellyfish as this species stings are not harmful to humans

(Dawson and Hamner 2005; Fautin and Fitt 1991). In contrast, the synergy of large

numbers of jellyfish and human usage of coastal environments has created several

problems, including power plant closures, challenges to fishery and aquaculture

operations (Purcell et al. 2007), and beach closures due to the pain and/or death of

beach-goers from jellyfish stings (Fenner and Williamson 1996).

Current data on jellyfish populations indicate that in certain regions of the world,

populations have increased (Condon et al. 2012), and two trends have been identified.

A weak trend since 1970 suggests that jellyfish have increased in relation to human

activities. Most notably are the increase in global temperature due to anthropogenic

production of carbon dioxide and overfishing. A strong trend over the last century

indicates that an oscillation in jellyfish populations may be a function of environmental

oscillations in ocean-atmosphere cycles and variations in insolation on food webs

(Condon et al. 2013). A recent study has documented the presence of jellyfish

3

populations along the majority of US coastlines, including Alaska and Hawai‘i, with

notable increases in the Northeast (Brotz et al. 2012). Thorough investigations into

jellyfish population dynamics require long term data sets, and these data are not

uniformly distributed worldwide (Condon et al. 2013; Hay 2006; Purcell et al. 2001). This

is especially true in estuaries, where jellyfish can be intermediate-top level predators

(Baird and Ulanowicz 1989; Condon and Steinberg 2008). As a consequence, the role

of jellyfish in food webs is often overgeneralized or misconstrued (Condon et al. 2012;

Purcell et al. 2007). Furthermore, because outreach education and associated social

media about scientific explorations stems from research (McKenna and Main 2013), it is

quite possible that the roles that jellyfish play in the environment are ambiguous at the

public interface. Jellyfish will continue to affect coastal communities as human usage of

coasts continues (Hinrichsen 1995) and jellyfish-human interactions are likely to

become more frequent. Thus, this research aims to provide perspective on the

interaction between jellyfish and at recreational areas.

North Carolina’s Albermarle-Pamlico estuarine system (APES) is the nation’s

second largest estuarine complex (Figure 1). APES provides important ecosystem

services, including essential habitat and nursery areas for a variety of east coast

fisheries and supports a substantial assortment of ecological, economic, recreational,

and aesthetic functions. APES was designated “an estuary of national significance” in

1987 (N.C. Department of Environmental and Natural Resources 2011).

Jellyfish are top predators in estuaries; however, studies on the distribution and

abundance of jellyfish within estuaries in general and APES in particular are limited. The

two most common jellyfish species are the scyphomedusan sea nettle, Chrysaora

4

quinquecirrha [Desor, 1848], and the ctenophore, Mnemiopsis leidyi [A. Agassiz, 1865],

(Miller 1974; Williams and Deubler 1968). The best record of these two jellyfish species

was noted in the Pamlico River Estuary (PRE) (Figure 1), where M. leidyi distribution

and abundance was documented by Williams and Deubler (1968), and spring/summer

abundance of the C. quinquecirrha (2 m-3) was calculated in 1967-68 by Miller (1974). A

study by Mallin (1991) on zooplankton distributions in the Neuse River Estuary (NRE)

reported the presence of jellyfish, but did not record species or abundance. Current

observations of the presence of other species of jellyfish, such as the cannonball

jellyfish Stomolophus meleagris [L. Agassiz, 1860], along the Outer Banks and Oak

Island (Figure 1) has been documented on-line by Appalachian State University

(http://www.jellyfish.appstate.edu/) and the Monterey Bay Aquarium

(http://jellywatch.org/) (Appalachian State University 2011; Elliott and Haddock 2010). It

should be noted that although the term “jellyfish” has been used to describe species

belonging to Phylum Cnidaria, Phylum Ctenophora and Phylum Chordata, Subphylum

Urochordata, Class Tunicata (Purcell 2012), unless otherwise indicated, I will use the

term “jellyfish” to describe species belonging to Phylum Cnidaria, Class Scyphozoa

(Figure 2).

I chose to investigate the potential drivers of jellyfish-human interactions in North

Carolina. My approach is described in three chapters: chapter 2 investigated physical

drivers of jellyfish-human interactions in the Neuse River Estuary (NRE) (Figure 1). The

NRE was selected as a study location for this chapter due to the annual presence of the

sea nettle, C. quinquecirrha, and because the NRE is a highly used recreational area. A

vital concern to NRE residents and stakeholders is recreation and tourism revenue, and

5

since sting of C. quinquecirrha are harmful to humans (Schultz and Cargo 1969), the

annual presence of C. quinquecirrha may have negative effects on NRE recreation and,

by association, tourism revenue. The Albermarle-Pamlico estuarine system (APES)

(Figure 1) and the NRE are greatly influenced by wind-driven circulation (Luettich et al.

2000; Reynolds-Fleming and Luettich 2004), and since C. quinquecirrha cannot swim

against water currents (Costello et al. 1998; Matanoski et al. 2001), I investigated how

wind affected the distribution and abundance of C. quinquecirrha in 2011 and 2012 at

six recreation sites located on the central north and south NRE shorelines. I used a null

hypothesis to test that the distribution and abundance of sea nettles would not differ at

all six recreation sites regardless of the wind dynamics, speed and direction. In addition

to analyzing wind, I also investigated other potential factors known to influence the life

history of the sea nettle and distribution, most notably salinity and temperature (Decker

et al. 2007; Calder 1974; Cargo and Schultz 1966).

The frequency of occurrence of jellyfish populations can also be directly

correlated to reproduction and growth (Hamner and Dawson 2008), and aggregations of

large numbers jellyfish or medusae may indicate sexual reproduction (Arai 1997). Thus,

the annual occurrences of C. quinquecirrha in the NRE may be undergoing sexual

reproduction. To evaluate the potential of sexual reproduction of C. quinquecirrha, I

used histology to determine gonad maturity, the presence of brooded larvae or

planulae, and the sex ratio of C. quinquecirrha collected throughout the 2011 field

season. The basis of my histological analysis stemmed from research done with C.

quinquecirrha in Chesapeake Bay where Littleford (1939) documented egg maturity, the

onset of fertilization, and development of planulae within female stomachs or gastric

6

cavities was documented. The size of jellyfish, which is typically measured by bell

diameter, may also influence spawning (Arai 1997) and in some jellyfish species bell

diameter may (Saucedo et al. 2012) or may not (Toyokawa et al. 2010) be linearly

related to egg diameter. I proposed several hypotheses: 1) the proportion of female and

male sea nettles would not differ, 2) measured egg diameters would be greater than

0.07 mm, indicating sexual maturity, 3) sexually mature males would have ruptured

sperm follicles or evidence of spent sperm follicles, 4) brooded planulae would be

observed in female gastric cavities indicating that sexual reproduction had occurred,

and 5) there would be no correlation between female C. quinquecirrha bell and egg

diameters. I also took the opportunity to compare visual observations of the sex of C.

quinquecirrha with my histology results since the color of mature gonads has been used

to classify sex (Littleford 1939).

The risk of receiving a sting by a jellyfish, like the sea nettle, is often a choice that

a coastal recreationist faces. The choice to recreate in water where jellyfish are clearly

visible is influenced by culture and human belief about jellyfish ecology. I chose to

evaluate cultural perspectives of jellyfish ecology and how jellyfish influence society

among four groups of people; fishers (recreation and commercial), recreationists

(surfers, swimmers, etc.), coastal researchers, and jellyfish researchers with a jellyfish

survey based on cultural consensus theory (CCT). CCT is a method used in

anthropology that allows researchers to quantify qualitative data, estimate cultural

beliefs, and to report an individual’s knowledge of those beliefs (Weller 2007). CCT is

based on culture or the set of learned beliefs, shared beliefs, and behaviors (Weller

2007). Cultural beliefs are affected by social norms or normative beliefs that are

7

associated with a group (Heywood 1996; Vaske and Whittaker 2004), and group culture

is the most frequently held items of knowledge and belief (D'Andrade 1987).

The cultural perspectives of jellyfish ecology among each group were assumed

to be different because knowledge of jellyfish in ecosystems is often misinterpreted

(Condon et al. 2012) and jellyfish are typically understudied despite being intermediate

to top level predators (Condon and Steinberg 2008). Therefore, it is likely that

misinterpretation of the ecological role of jellyfish would extend to the public interface

because most outreach education and associated social media about scientific

explorations stems from research (McKenna and Main 2013). To test ecological literacy

concerning jellyfish, I hypothesized that the cultural perspectives of jellyfish ecology

would not differ among fishers, recreationists, coastal researchers, and jellyfish

researchers. Jellyfish-human interactions may depend on jellyfish species and

specifically if the jellyfish stings are harmful to humans. Often jellyfish are viewed as

“nuisance” species (Richardson et al. 2009) and media reports about jellyfish can be

negative (Condon et al. 2012). I tested the null hypothesis that all groups would share

similar cultural perspectives of jellyfish.

The goal of my dissertation research was to add to the growing knowledge of

jellyfish research by studying jellyfish-human interactions in North Carolina. Specifically,

my results will indicate: 1) if jellyfish interactions at recreational sites can be related to

abiotic conditions. Such an outcome would indicate that periods of high jellyfish

encounter rate at recreational sites may be predictable; 2) if sexual reproduction is

occurring at different recreational sites during jellyfish aggregation events. If sexual

reproduction is occurring it would indicate that management of jellyfish in the APES

8

system should be focused on the medusa stage; and 3) if there are particular areas of

human perception of jellyfish ecology that may be the focus of educational outreach and

if particular characteristics of jellyfish perception that may be targets of regulation. To

date, multifaceted research approaches similar to that which I have adopted have

helped manage encounters with jellyfish species in Australia (Gershwin et al. 2010) and

Germany (Baumann and Schernewski 2012). To understand the global extent of

jellyfish ecological and socio-economic implications, the National Center for Ecological

Analysis and Synthesis (NCEAS) developed the “Global expansion of jellyfish blooms”

working group, to analyze and synthesize existing jellyfish data. With more data

available on jellyfish in coastal environments, the management and sustainability of

coastal resources by local, state, and federal agencies will improve in areas prone to

jellyfish-human interactions.

9

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Decker, M. B., C. Brown, R. Hood, J. Purcell, T. Gross, J. Matanoski, R. Bannon & E.

Setzler-Hamilton, 2007. Predicting the distribution of the schyphomedusa

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14

LIST OF FIGURES

Figure 1. Map of eastern North Carolina, including the Albermale Pamlico Estuarine

System; Albermarle Sound, Pamlico Sound, Pamlico River Estuary (PRE), and the

Neuse River Estuary (NRE).

Figure 2. Conceptual diagram of the scyphozoa life cycle based on the life history of the

sea nettle (C. quinquecirrha). Sexual reproduction of medusae yields a zygote that will

metamorphosize into a larva or planula. The planula will swim toward the bottom of the

estuary and settle on a substrate (i.e. rocks or oyster shells) and turns into a polyp,

which is also known as a scyphistoma (Cargo and Schultz 1966). The polyp will

reproduce asexually by budding (B), pedal laceration (C), and/or transverse fission.

Unique to jellyfish reproduction is a type of transverse fission called strobilation (Arai

1997), where the polyp will develop into a strobila and planktonic jellyfish or ephyrae are

produced when temperatures are elevated (Calder 1974). The ephyrae will mature into

medusae and the life cycle repeats (Arai 1997).

15

Figure 1.

16

Figure 2.

CHAPTER 2

Local wind dynamics influences the distribution and abundance of the sea nettle,

Chrysaora quinquecirrha, at six estuarine recreation sites in the Neuse River Estuary

18

ABSTRACT

The “stinging” sea nettle, Chrysaora quinquecirrha, has been observed annually

in the Neuse River Estuary (NRE), a site of intense human recreation in North Carolina.

The purpose of this chapter was to evaluate what physical drivers influence jellyfish-

human interactions. Jellyfish were counted and abiotic variables were measured bi-

weekly at six recreation sites from May to August, 2011 and 2012. In particular, the

influence of wind on jellyfish abundance was investigated as circulation within the NRE

is primarily a function of wind. The two years differed in stream flow, salinity, and

temperature, due to drought conditions in 2011. Wind speed and direction did not differ

between the two years; however, the number of high-speed wind events did differ

across years at the Cherry Point Marine Weather Station. A total of 3,241 jellyfish were

observed, with peak counts in July and minimum counts in August. There were

significant differences in mean sea nettle counts between sites, specifically between

north and south shorelines. Northeast-east and southeast-southwest wind events (3 – 8

m s-1) measured from Cherry Point and SW wind events (2 – 7 m s-1) measured from

Cape Hatteras were correlated to sea nettle abundance (m2) when a lag period was

included. I conclude that NRE wind dynamics are one of the factors influencing jellyfish-

human interactions at the six recreational sites in the NRE. The relatively strong

influence of wind, compared to the abiotic variables temperature and salinity, suggests

that prediction of sea nettle abundances at these sites subsequent to wind events is

possible.

19

INTRODUCTION

Large numbers of cnidarian scyphomedusae (jellyfish) have created problems in

coastal environments (Brotz et al. 2012; Condon et al. 2011; Mills 2001; Purcell 2012;

Purcell et al. 2007). These deleterious effects include power outages due to clogged

cooling intake-valves of power plants (Matsueda 1969; Matsumura et al. 2005;

Rajagopal et al. 1989; Yasuda 1988), severe damage to commercial fishing and

aquaculture operations due to net bursting (Graham et al. 2003; Nagata et al. 2009),

tainted catches and predation of farmed fish (Purcell et al. 2007), and beach closures

and injury (including death) to coastal recreationists (Fenner and Williamson 1996;

Pages 2001). Coastal and estuarine environments are conducive for jellyfish blooms in

that they are characterized by seasonal pulses of nutrients, ample habitat for

benthic/juvenile phases of jellyfish, and an environment to aggregate for reproduction

(Lo et al. 2008; Omori and Nakano 2001; Pitt and Kingsford 2000; Purcell 2005; Purcell

2012). Therefore, when compared to the open ocean, coastal and estuarine jellyfish

species are observed in higher densities and may have seasonal mass occurrences

(Decker et al. 2007; Doyle et al. 2007; Hamner and Dawson 2009).

High abundances of the sea nettle Chrysaora quinquecirrha [Desor, 1848] have

been reported in the Neuse River Estuary, North Carolina, during the spring and

summer (Mallin 1991) (Figure 1A). Chrysaora quinquecirrha is a brackish water (salinity

5 – 18) jellyfish species (Cargo and Schultz 1966) that is highly abundant during the

spring and summer months in mid-Atlantic estuaries, e.g. Chesapeake Bay (Virginia

and Maryland) (Calder 1974; Calder 1972; Cargo and Rabenold 1980; Cargo and King

1990; Decker et al. 2007; Mansueti 1963), the Albermarle-Pamlico Estuarine System

20

(North Carolina) (Mallin 1991; Miller 1974; Williams and Deubler 1968), and Barnegat

Bay (New Jersey), USA. The most common negative interaction between jellyfish and

humans is a sting (Purcell et al. 2007), and although all species belonging to Phylum

Cnidaria have stinging cells or nematocysts, the extent to which a jellyfish sting will

affect humans is variable from “feeling nothing” to life threatening (Arai 1997; Gershwin

et al. 2010; Mills 2001). The pain associated with sea nettle stings have been described

as “bothersome” to “extremely painful” (Cargo and Schultz 1966; Decker et al. 2007). To

circumvent harm to humans, barrier nets have been utilized in swimming areas due to

the presence of C. quinquecirrha in Chesapeake Bay (Schultz and Cargo 1969) and

Barnegat Bay, New Jersey. It is unknown how sea nettles affect coastal tourism and/or

recreation within the NRE (Figure 1B) and the greater APES, but an assessment of

NRE stakeholder interests revealed that recreation and maintaining tourism income

were important public interests (Maloney et al. 2000). Coastal recreationists in the NRE

desired to be safe in the water when engaging in prolonged-body contact activities (i.e.

swimming and water skiing) and ‘getting rid of “slime” on fishing nets (Maloney et al.

2000). Although “slime” was not properly defined by Maloney et al. (2000), jellyfish are

known to clog fishing nets and in consequence the gelatinous bodies of jellyfish are

often torn apart (Nagata et al. 2009; Purcell 2009) and could resemble “slime.”

Similar to other reports of jellyfish occurrences, the presence or absence of C.

quinquecirrha along NRE shorelines has been considered ephemeral and

unpredictable. It is plausible that the combination of jellyfish swimming behavior, wind

velocity and local hydrology, topography, and bathymetry contribute to the advection of

jellyfish within coastal areas and in some cases an increase in abundance (Cargo and

21

King 1990). Chrysaora quinquecirrha have been described as “cruising predators,” that

will constantly swim in order to maximize prey capture via water vortices generated by

bell contractions or pulsation (Ford et al. 1997). Although pulsation rate and swimming

velocities will increase when prey is present, most of the swimming directions are

vertical rather than horizontal in the water column and maximum swimming velocities

are approximately 1.8 cm s-1 (Costello et al. 1998; Matanoski et al. 2001). Furthermore,

in situ observations of C. quinquecirrha swimming toward or into tidal currents did not

achieve overall forward direction (Costello et al. 1998). On average, water current

velocities measured along and across the NRE are approximately 5-10 cm s-1 (Luettich

et al. 2000).

Pamlico Sound (Figure 1A) and NRE estuarine circulation is dominated by wind

and density currents due to the large volume of shallow water that encompasses this

system and negligible tidal ranges (0.15 m near the mouth and 0.3 m in the upper

estuary) (Luettich et al. 2000; Pietrafesa et al. 1986; Roelofs and Bumpus 1953; Stanley

and Nixon 1992). Wind blows across Pamlico Sound and the NRE from the south (S) to

southwest (SW) between April and August and from the northwest (NW) to northeast

(NE) between September and February (Wells and Kim 1989). The presence of a wind-

tide (13.2 hr period) has been documented in the NRE due to a wind-induced seiche

within Pamlico Sound (Luettich et al. 2002). Depending on wind magnitude, it takes

roughly one to two weeks for water to move in and out of the NRE to Pamlico Sound.

Water is pushed into the NRE from Pamlico Sound with winds originating from the NE

and conversely, water is driven out of the NRE with SW winds (Luettich et al. 2002;

Luettich et al. 2000). Water currents move across the channel of the NRE with SW

22

winds moving water toward the north shoreline and NE winds moving water toward the

south shoreline. These across-channel circulation dynamics also influence salinity and

water level (Reynolds-Fleming and Luettich 2004). Observations of along and across-

channel circulation in the NRE may vary from within a day to about two weeks (Luettich

et al. 2002; Luettich et al. 2000; Reynolds-Fleming and Luettich 2004) and this results in

a delayed observed response in the presence or absence and abundance of sea nettles

along the NRE coast. A geomorphologic characteristic of the NRE is its distinct “V”

shape (Figure 1) with the upstream section oriented NW-SE and the downstream

section oriented SW-NE. This orientation, coupled with shallow water and dominant

wind directions from the NE and SW creates a significant vector for longitudinal wind

forcing in the NRE (Reynolds-Fleming and Luettich 2004). The sharp bend in the middle

of the NRE has been used as a midway marker in many North Carolina estuarine

studies (Buzzelli et al. 2002; Luettich et al. 2000; Reynolds-Fleming and Luettich 2004).

This midway area is also the narrow section of the river estuary, which crosses the

Minnesott sand ridge; a feature associated with the regional Suffolk Shoreline, a

stranded Pleistocene paleo-shoreline feature (Mallinson et al. 2008; Parham et al.

2013). Since water is moved along and across this narrow NRE bend (Luettich et al.

2000), this area could serve as an accumulation area for C. quinquecirrha, especially if

wind-generated water currents exceed the threshold of C. quinquecirrha swimming

capabilities.

The life history of C. quinquecirrha and how abiotic variables influence the

biology of this species has been thoroughly studied in Chesapeake Bay. Preliminary

work done by Cargo and Schultz (1966), Calder (1972, 1974), and Cargo and Rabenold

23

(1980) documented the development and behavior of the polyp, strobila, and ephyrae in

both laboratory and field observations. An ecological forecasting system has been

developed to predict the distribution of medusae within Chesapeake Bay based on

temperature (26-30°C) and salinity (10-14) (Decker et al. 2007). Within large bays and

estuaries, including Chesapeake Bay, C. quinquecirrha are dominant carnivores that

are adapted to estuarine salinities (Decker et al. 2007; Hamner and Dawson 2009;

Purcell and Decker 2005).

The purpose of this chapter was to investigate if physical factors could be

correlated to the distribution and abundance of NRE C. quinquecirrha at six recreation

sites (Figure 1B). Since the NRE is heavily influenced by wind-driven circulation, has a

geomorphology conducive for the possibility of jellyfish accumulation, and C.

quinquecirrha are weak swimmers, I tested the null hypothesis that the distribution and

abundance of C. quinquecirrha would not differ at NRE six recreation sites. I also

investigated whether other abiotic variables could be correlated with the distribution and

abundance of this species at these six sites (Figure 1B). The overall objective of this

chapter was to determine if jellyfish-human interactions at the six recreational sites may

be influenced by local estuarine conditions. If this is the case, it may be possible to

predict when jellyfish abundance may increase at these sites.

MATERIALS AND METHODS

Study area and recreation sites

The Neuse River Estuary (NRE) is a shallow estuary located SW of Pamlico

Sound, North Carolina (Luettich et al. 2000) (Figure 1). It occupies a drowned river

24

valley that flooded and filled during the last post-glacial sea-level rise, beginning

approximately 18,000 years ago (Wells and Kim 1989). The length of the NRE is

approximately 70 km, with a mean width of 6.5 km and mean depth of 3.6 m (Luettich et

al. 2000). For the purposes of this chapter, the estuarine shorelines of the NRE will be

referred to as “north” and “south” shorelines (Figure 1B), with the horizontal axis located

at the center of the NRE bend. Each shoreline had three recreation sites that spanned a

horizontal distance of 15 km. These research sites were selected due to the high

number of sea nettles observed annually by coastal recreationists and all sites are

highly used for recreation each spring and summer.

The north shoreline sites included the Town of Oriental’s marina and two YMCA

camps called Camp Sea Gull and Camp Seafarer. Oriental (OM) is located along the

north bank of the NRE (Figure 1B). It is a town that has more registered boats (~1,200)

than residents (~825) and offers a range of coastal activities, including sailing, fishing,

kayaking, wind surfing, etc. for residents and tourists. Camp Sea Gull (CSG) and Camp

Seafarer (CSF) (Figure 1B) provide activities/programs for children, teens, and adults

through spring, summer, and fall. Each camp spans 8 km (5 mi) along the NRE, with

approximately 0.9 km (3000 ft) of estuarine shoreline for seamanship and other water

activities. During the summer camp session, each camp hosts approximately 800

campers (age 6-16).

The south shoreline sites surveyed belong to the “Neuse River Recreation Area”

of the USDA Croatan National Forest; a federal national forest that encompasses

651.54 km2 (161,000 acres) of land and offers a vast array of recreational activities for

the general public. I chose three sites within this area: Flanners Beach (FB), Pine Cliff

25

(PC), and Siddie Fields (SF) (Figure 1B). All three sites were used as research sites

because each site has an extensive beach area for recreation and frequent

observations of sea nettles and therefore, a greater potential for jellyfish-human

interactions.

Sea nettle abundance and in situ abiotic data collection

The abundance of C. quinquecirrha was surveyed using visual counts at each

site within a 12-hr diurnal period, twice a week, from spring (Mid-May) to summer (Mid-

August), for two consecutive years (2011-2012). On each observation day, two to four

researchers counted C. quinquecirrha and all observations were conducted in unison. In

2011, observations did not begin until 1 June, but C. quinquecirrha were noted in Mid-

May. In 2012, observations started in Mid-May. On each observation day, there were at

least two researchers that conducted visual counts and C. quinquecirrha were surveyed

at two types of coastal-recreation interfaces, a coastline and/or pier. At the coastline

research sites (FB, PC, and SF) a 100 m transect was followed along the water’s edge

and jellyfish in the water or washed ashore were counted along the transect line, in

adjacent coastal waters 2 m from the transect line, and in water depth less than 1 m. At

CSG and CSF, coastline and pier counts were conducted. Coastline counts at each

camp site were performed in the same manner as the Neuse River recreation area. A

pier count was also performed at the Town of Oriental’s marina (OM). All pier counts

included C. quinquecirrha within a 2 m distance from the pier.

To standardize researcher effort and extent of observations, a 1-sec footstep

pace was employed at the start and end of the transect line and/or pier. It should be

26

noted that unlike the coastline surveys, each pier length was variable among research

sites; Camp Seafarer (~200 m), Camp Sea Gull (~280 m), and Oriental Town Marina

(~215 m). However, since all observations were conducted at a 1-sec footstep pace, the

time spent observing jellyfish was relative to pier length. It should be noted that for all

statistical analyses, C. quinquecirrha counts were standardized to 100 m length

observations to assure that the amount of observation time was consistent among

research sites. At each site, a Conductivity Temperature Depth meter (YSI CastAway)

was used to measure temperature (ºC), salinity, and depth (m). Secchi disk (m) was

used to approximate water turbidity. These measurements were collected from midway

markers at each coastline site and in the middle and end of all pier sites. Daily stream

flow (m3 s-1) data were collected from the USGS Fort Barnwell location. These abiotic

variables were selected for this study because C. quinquecirrha medusae are greatly

influenced by these variables in Chesapeake Bay (see Cargo and Schultz 1966, Cargo

and King 1990, and Decker et al. 2007).

Statistical analyses and data acquisition

Sea nettle abundance (m2) was tabulated for each site and observation day to

investigate how abundance differed by month and site, and the standard deviations of

all counts were calculated to account for variability in researcher observations. If the site

had coastline and pier interfaces, the interface with the greatest number of C.

quinquecirrha was used. Since C. quinquecirrha abundance did not fit a normal

distribution, the data were transformed (Log+1) in order to fulfill the requirements of the

parametric tests. Variation in mean sea nettle count was then assessed between sites,

27

months and year using a three-way ANOVA. For this analysis, since no data were

collected in May 2011, only the months June, July, and August were used for both

years. To compare annual mean C. quinquecirrha counts (Log+1) in 2011 and 2012, a t-

test was performed. Comparisons were also made for abiotic variables between years

using t-tests.

Pearson correlations were used to compare C. quinquecirrha abundance and all

abiotic variables by year and by shoreline. Wind data were acquired from the National

Weather Service, NOAA weather station at Cherry Point Marine Base and Cape

Hatteras Weather Station, North Carolina. These weather stations were the same used

by Luettich et al. (2000, 2002) and Reynolds-Fleming and Luettich (2004) to generate

models of wind-driven circulation in APES, the NRE and the Renaissance Computing

Institute (RENCI), Coastal Emergency Risks Assessment (NC-CERA), ADCIRC Coastal

Circulation and Storm Surge Model + SWAN Wave Model. Wind variables included daily

wind speeds (m s-1) and directions (North = 0°) that were calculated from hourly data.

To observe wind dynamics, wind diagrams were created to observe daily changes in

wind speed (m s-1) and direction (North = 0°). Pearson correlations were used to

determine if wind events (when wind speed increased by at least 4 m s-1 over the

course of 6 hours) were correlated to C. quinquecirrha abundance and distribution. The

time frame of a week prior to observations was used for wind speed, wind direction, and

wind event analyses because variable wind speeds and directions affect surface current

vectors in the NRE within this time frame (Luettich et al. 2000; Luettich et al. 2002). To

help create conceptual diagrams of C. quinquecirrha and wind dynamics, NRE water

velocity and direction data were acquired from model runs generated by the RENCI NC-

28

CERA ADCIRC model from nodes within 100 – 300 m of FB in 2011 and all research

sites in 2012.

RESULTS

Abiotic variables compared by year

Mean temperature, salinity, stream flow, and wind speed measured at Cherry

Point were all significantly different in 2011 and 2012 (Table 1). The range in stream

flow and salinity was considerably smaller in 2011 than 2012 (Table 1, Figure 2). There

was also a larger range in wind speed in 2011 (Figure 3). Mean wind directions

measured at Cherry Point as well as mean wind speed and direction measured from the

Hatteras weather station were not significantly different (Figure 3).

Variation in C. quinquecirrha abundance

A total of 3,241 ± 800 (standard deviation) C. quinquecirrha were counted at all

research sites in 2011 and 2012 (Figure 4). The t-test showed that mean C.

quinquecirrha counts for 2011 and 2012 were not significantly different (t = -0.47, DF =

729.3, p-value = 0.64). Overall, the majority of C. quinquecirrha were counted in July

and the least in August (Figure 4). The three-way ANOVA revealed significant

differences in mean C. quinquecirrha count by sites and months and significant

interactions among all three factors (Table 2).

Abiotic parameter and wind correlations

29

Most of the C. quinquecirrha abundance correlations with in situ abiotic variables

were not significant on the north (Table 3) and south (Table 4) shorelines. Weak but

statistically significant correlations were noted with stream flow on the north shoreline (R

= -0.20, p-value = 0.01) and depth (R = -0.19, p-value = 0.02), secchi (R = -0.19, p-

value = 0.02), and salinity (R = 0.18, p-value = 0.04) on the south shoreline. Wind

speeds measured from Cherry Point one and two days prior to observations and wind

data measured from Hatteras also had weak correlations with wind direction two days

prior and wind speeds one and two days prior to north shoreline observations (Table 3).

On the south shoreline, wind speed measured from Cherry Point one day prior to

observations was correlated with C. quinquecirrha abundance. Significant correlations

were also noted with wind direction and speeds measured from Hatteras one and two

days prior to observations (Table 4).

Wind events and C. quinquecirrha abundance

Wind event correlations were higher than the correlations with daily averaged

wind speeds and directions (Table 3 and Table 4). During this study, wind events of 3 to

8 ms-1 (N = 7) measured at Cherry Point occurred in July 2011, May 2012, and June

2012 (Figure 6) and correlations were observed with wind direction five days prior to

observations on the north shoreline (Table 5) and one day prior to observations on the

south shoreline (Table 6). Wind events from 2 to 7 ms-1 (N = 8) measured at Hatteras

occurred in May, June 2011 and May, June and July 2012 (Figure 7) and there were no

significant correlations made with abundance on the north shoreline. A negative

correlation (R = -0.54, p-value = 0.02, Table 6) with C. quinquecirrha abundance on the

30

south shoreline and wind direction two days prior to observations was observed. It

should be noted that wind events up to 6 ms-1 were observed with the greatest

frequency (Figure 6 and Figure 7). When wind events of 3 to 8 ms-1 were measured

from Cherry Point, C. quinquecirrha abundance ranged from 50 to 280 m2 on the north

and south shorelines with wind directions SE, S, and SW (Figure 8A). A smaller range

of abundance measuring 25 to 75 m2 was associated with NE and E wind directions and

in general, larger abundances were recorded on the south shoreline (Figure 8B). The

distribution of C. quinquecirrha along the south shoreline was not uniform, instead,

higher abundances were observed at PC and SF when wind events occurred from a

SSE to a SSW direction (Figure 8B). Wind events in August were not included in these

analyses because no jellyfish were observed in August 2011 and < 10 jellyfish were

observed in August 2012.

DISCUSSION

Wind dynamics and C. quinquecirrha in the NRE

My data showed that wind speed and direction are correlated to the distribution

and abundance of C. quinquecirrha in the NRE; therefore, I rejected my null hypothesis

that there is no difference in sea nettle abundance among the six recreational sites. It

was interesting that wind dynamics appear to be more influential to NRE jellyfish-human

interactions than other abiotic variables, including temperature and salinity. When wind

events occurred, I found that SSE to SSW wind directions one day prior to observations

could be correlated with increased numbers of C. quinquecirrha along the south

shoreline, particularly at stations PC and SF (Figure 1, Figure 8B). The highest

31

abundances of 300 m2 observed at PC and SF were correlated to S to SW winds

(Figure 8B) and because C. quinquecirrha are weak swimmers with maximum

swimming velocities of 1.8 cm s-1 (Costello et al. 1998; Matanoski et al. 2001) it is likely

that these abundances are a function of the water currents, shallow waters, and

geomorphology of the NRE.

Wind dynamics and stream flow are two major factors that influence how water

moves in the NRE (Luettich et al. 2000) and in 2011 there was significantly less stream

flow than in 2012 (Table 1, Figure 2). Weak stream flow and averaged water velocities

obtained from ADCIRC data near Flanners Beach (Figure 1B) showed along-channel

water movement (currents toward the NW and SE) instead of across-channel water

movement (currents toward NE and SW) as observed in 2012 (RENCI 2012). However,

with no data available for the other sites in 2011, I cannot conclude that water velocities

and direction were similar or an anomaly of the 2012 data. Therefore, although water

directions observed near Flanners Beach may have been different in 2011 and 2012, I

do not attribute this variation to an increase or decrease in C. quinquecirrha abundance

(Figure 4). Instead, understanding that during June and July, wind speed and direction

are the primary drivers of water currents that move either along or across the channel of

the NRE (Luettich et al. 2000; Reynolds-Fleming and Luettich 2004); I suggest that

variation in C. quinquecirrha abundance is a function of wind-driven circulation more so

than stream flow dynamics. Surface currents of 5 to 13 cm s-1 will move water

downstream (east of the bend) when SW winds were measured at 8 m s-1 (Luettich et

al. 2000; RENCI 2012). Movement of jellyfish is likely to follow the downstream direction

of the surface current and accumulation occurs at the bend of the NRE, which coincides

32

with the PC and SF locations (Figure 9A) and water current directions obtained from the

RENCI ADCIRC model.

Downstream surface currents generated from SW winds will also decrease in

speed when winds shift to opposing directions and this can occur in short periods of

time from hours to within a week (Luettich et al. 2000). These downstream surface

current dynamics could explain why large numbers of C. quinquecirrha were not

uniformly observed throughout the field seasons at PC and SF. All south shoreline sites

were coastline interfaces with beaches and large expanses of shallow water, therefore,

it is plausible that C. quinquecirrha were washed ashore by surface currents and

confined to these areas with shallow water depths. Water is also moved into NRE from

Pamlico Sound via a NE wind; and when wind events were measured from Cherry

Point, the movement of water toward the south shoreline could also favor the

accumulation of C. quinquecirrha at the bend of the NRE (Figure 9B). A SW wind could

also move C. quinquecirrha away from this accumulation area, which is near PC and

SF, depending on water currents near the bend of the NRE and this may explain why I

noted a negative correlation (R = -0.54, p-value = 0.02) with wind events measured

from Hatteras two days prior to observations (Figure 9C). These observations may

reflect the 13.2 hr wind tide, or seiche of water that moves in and out the NRE with NE

and SW winds (Luettich et al. 2000).

Water depth is also influenced by water currents that move across the NRE

channel; SW winds 2 to 8 m s-1 will lower water depths on the south shoreline

(Reynolds-Fleming and Luettich 2004) and this contributed to an increase in south

shoreline C. quinquecirrha abundance (Figure 9D). South shoreline C. quinquecirrha

33

abundance was also correlated with NE wind events and because NE winds 2 to 5 m s-1

generate currents 5 to 10 cm s-1 that move toward the south shoreline (Luettich et al.

2000), cross channel water circulation may also explain accumulations of jellyfish on the

south shoreline via wind (Figure 9E). Although the wind events correlation (R = 0.53, p-

value = 0.00) was weaker on the north shoreline versus the south shoreline, NE winds

can upwell bottom water along the north shoreline, which could bring C. quinquecirrha

to the surface and explain why jellyfish abundance also corresponded to NE wind

events five days prior to observations (Figure 9E). A five day lag in C. quinquecirrha

observations reflected the delayed response of water movement with wind dynamics as

described by Luettich et al. 2000 and Reynolds-Fleming and Luettich 2004 in the NRE.

Additionally, SW winds 7 m s-1 can generate water currents up to 5 cm s-1, which move

water toward the north shoreline (Luettich et al. 2000), increasing the likelihood of more

C. quinquecirrha being pushed toward the north shoreline (Figure 9D). It should be

noted that lower abundance was consistently observed at the north shoreline

recreations sites (Figure 8A). A possible explanation for this is all north shoreline sites

were pier interfaces, which made accumulation and confinement of jellyfish less likely to

occur at these recreation sites. Therefore, future abundance studies should use beach

seines and trawl nets to compare if there are significant differences in sea nettle

abundance on the south and north shorelines, respectively.

C. quinquecirrha life history stages and inference to North Carolina

I did not find significant correlations with C. quinquecirrha medusa abundance

and temperature and/or salinity at the six NRE recreation sites. This was surprising as

34

these variables correlate strongly with medusa presence in Chesapeake Bay (Decker et

al. 2007). It is possible that the shallow nature of the NRE yields different relationships

with the abiotic variables commonly associated with life history stages of C.

quinquecirrha. The NRE can be a strongly stratified estuary or a well-mixed estuary

depending on wind direction (Luettich et al. 2000), therefore temperature and salinity

correlations with C. quinquecirrha may be similar to Chesapeake Bay observations but

at shorter time intervals. Field research in Chesapeake Bay noted that ephyrae

production started in April and continued to early July; however, the highest number of

ephyrae produced was during the spring (Calder 1974; Cargo and Rabenold 1980).

Ephyrae of C. quinquecirrha have not been documented in North Carolina; however, the

movement of ephyrae in the NRE can be inferred using Chesapeake Bay life history

and APES estuarine circulation literature. In the NRE, Wells and Kim (1989) reported

that the highest freshwater input occurred in February (high precipitation, low

evaporation) and the lowest freshwater input occurred in June (low precipitation, high

evaporation). Therefore it is likely that the dominant circulation type, estuarine

circulation, is prominent during February and extends into spring. Calder (1974) noted

that ephyrae production is highest in the spring in Chesapeake Bay and when dispersed

ephyrae will swim toward bottom waters (Calder 1974). This behavior coupled with the

estuarine circulation noted during spring months, would favor the movement of ephyrae

into the NRE from Pamlico Sound via estuarine circulation and these ephyrae would be

isolated therein.

Ephyrae production occurs at a bi-monthly rate (Calder 1974); therefore

aggregations of C. quinquecirrha medusae in APES and the NRE should be similar to

35

Chesapeake Bay where jellyfish are observed in variable frequencies and abundances.

With adequate nutrition, ephyrae can grow into medusae within a month (Calder 1972;

Olesen et al. 1996). Therefore, based on Chesapeake Bay literature and NRE estuarine

circulation dynamics, the C. quinquecirrha medusa stage should be present toward the

end of spring, with highest numbers in the summer (Calder 1972; Decker et al. 2007). In

our two year study, C. quinquecirrha presence began in Mid-May and decline began in

late July at the six recreation sites in the NRE (Figure 1B). These observations coincide

with Miller’s (1974) C. quinquecirrha observations conducted in the PRE (Figure 1A).

Miller (1974) attributed the spring presence of C. quinquecirrha in the PRE to dramatic

decreases in M. leidyi abundance and noted that M. leidyi abundance steadily declined

until the end of July or as long as C. quinquecirrha were present. It is well known that C.

quinquecirrha feed on M. leidyi and this relationship has been thoroughly documented in

many Chesapeake Bay food web studies and experiments (Baird and Ulanowicz 1989;

Cargo and Schultz 1967; Feigenbaum and Kelly 1984; Purcell 1992; Purcell et al.

1994a; Purcell et al. 1994b). I observed M. leidyi in the early spring of each field season

and made note that by the end of June none were present at the study sites. Other than

these observations, no surface counts and/or abundance surveys were taken of M.

leidyi. To date, C. quinquecirrha abundances in the PRE have been reported to be 2 m-3

via net sampling (Miller 1974).

It is also possible that C. quinquecirrha polyps have an established habitat within

the NRE and ephyrae are moved around the NRE by wind-induced water circulation.

Possible NRE polyp habitat sources include docks or oyster sanctuaries (Figure 1A)

and C. quinquecirrha polyps appear to preferentially settle on oyster shells (Breitburg

36

and Fulford 2006; Calder 1974). Since 1996, the North Carolina Department of

Environment and Natural Resources, Division of Marine Fisheries (DMF) has launched

an oyster sanctuary program where ten oyster sanctuaries have been established within

Pamlico Sound (Figure 1A), with a sanctuary located in the NRE (Figure 9), to develop

and protect native oyster brood stock and increase biodiversity. If these oyster

sanctuaries provide habitat for C. quinquecirrha polyps, then they are a likely source

population for medusae to enter the NRE. In contrast, it is also quite possible that the

oyster sanctuaries may not be suitable habitat for C. quinquecirrha polyp populations.

To date, no in situ experiments have evaluated this potential nor the contribution and

possible preference of human supplied substrate for C. quinquecirrha polyps in North

Carolina Estuaries, including the NRE.

In this study, C. quinquecirrha abundance was calculated from surface counts

divided by the area of observation (100m length of coastline or pier * the field of view

per observer 2m) multiplied by the depth of observations (1m). From this calculation, we

concluded that C. quinquecirrha abundance varied from 0 to ~2 per m3, with greater C.

quinquecirrha abundances noted at the NRE south shoreline sites in July (Figure 9).

Although different methods were employed in this study and Miller’s study in 1974, the 2

m-3 C. quinquecirrha abundance observations and the seasonal presence/absence of C.

quinquecirrha noted raises two interesting points about the C. quinquecirrha populations

in North Carolina. First, the abundance of C. quinquecirrha populations in North

Carolina seems to be much less than those sampled in Chesapeake Bay, where the

highest abundance of ~16 m-3 occurs in July and August (Purcell 1992). Second, the

sea nettle season in North Carolina reduces to a few sighted jellyfish in August whereas

37

sea nettles in Chesapeake Bay are present in low abundances through the fall until

October (Purcell et al. 1994b). Longer term studies on the biology of C. quinquecirrha

are needed to confirm these abundance observations and accurately report sea nettle

population dynamics in North Carolina.

CONCLUSIONS

My study has shown that jellyfish-human interactions in the NRE are influenced

by wind speed, wind direction, and subsequently water current direction. Prevalent SE

to SW wind events cause large numbers of C. quinquecirrha to accumulate at recreation

sites on the south shoreline one day after a wind event and significant increases or

decreases may be noted on the north shoreline five days after a wind event, depending

on direction. Rosa et al. (2012), also correlated wind speed and SE direction to an

increase in the abundance of Pelagia noctiluca [Fosskaal, 1775] (R = 0.2, p-value =

0.01) at a sheltered research site (S. Agata) in the Straits of Messina, Italy. However,

unlike my study where wind was a very influential factor in C. quinquecirrha abundance,

stronger correlations were made with temperature and the abundance of P. noctiluca

(Rosa et al. 2012). In addition to wind direction, it is known that Langmuir circulation is

associated with patch formations of jellyfish species (Hamner and Schneider 1986;

Larson 1992). In Belize, Larson (1992) observed Linuche unguiculata [Roberts, 1827] in

slicks and windrows with flotsam that are oriented parallel to wind, which are indications

of Langmuir circulation. The L. unguiculata are aggregated at the convergence zones

between adjacent Langmuir cells and remain there by the jellyfish’s upward swimming

behavior. Circular swimming behavior also reduces the rate of patch dispersion and

38

may explain why jellyfish patches were observed when Langmuir circulation was not

prominent in March and April (Larson 1992). In the Bering Sea, species of

hydromedusae and scyphomedusae are also aggregated at surface waters between

Langmuir convergence cells (Hamner and Schneider 1986). These aggregations form

rows between the areas of convergence and divergence that may be evenly dispersed

by 100 m. This linear pattern may be advantageous for jellyfish predation, especially at

night when planktonic prey will migrate vertically to surface waters where the jellyfish

are gathered. However, confinement of jellyfish to the surface waters also increases the

likelihood of predation, especially from seabirds (Hamner and Schneider 1986).

Wind dynamics may be used to potentially predict the distribution and abundance

of C. quinquecirrha in the NRE; therefore, these data may be used to reduce harmful

jellyfish-human interactions and thereby reducing the possibility of loss in

tourism/recreation revenue. Jellyfish stings in tourism and/or recreations areas have

been an on-going coastal management problem globally and coastal communities have

responded by attempting to minimize these jellyfish-human interactions. For example, in

Australia, public awareness, mitigation devices, and on-site first-aid stations have been

made available to tourists (Gershwin et al. 2010). Public education and awareness,

including information posted on signs and pamphlets about jellyfish given to beach-

goers, has also helped people cope with large numbers of jellyfish on German beaches

(Baumann and Schernewski 2012). In Chesapeake Bay, large numbers of the C.

quinquecirrha have affected coastal recreationists since the 1960’s (Cargo and Schultz

1966). Fortunately, over 50 years of research on this stinging jellyfish species in

Chesapeake Bay has provided the foundation for a monitoring system, which has

39

continued to accurately predict the probability of encountering a sea nettle (Decker et al.

2007). In areas where extensive jellyfish research is limited, public monitoring of high

numbers of stinging species has been conducted to reduce harmful jellyfish-human

interactions. Examples of these observations include the high likelihood of occurrence

of the harmful box jellyfish species historically referred to as Carybdea alata [Reynaud,

1830] but currently regarded as Alatina moseri [Mayer, 1906] (Bentlage et al. 2010;

Gershwin 2005) on Waikīkī Beach, Hawai‘i, 8 to 12 days after a full moon (Thomas et

al. 2001), and more recently C. quinquecirrha abundances are frequently reported to the

public by the Barnegat Bay Partnership in Barnegat Bay, New Jersey. The use of data

gathered from research and public interfaces will increase the likelihood of successful

management of jellyfish-human interactions.

40

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48

LIST OF TABLES AND FIGURES

Table 1. T-test comparison of abiotic variables by year; N = 139. Depth, secchi,

temperature and salinity variables were collected in situ. Stream flow data was obtained

from the USGS Fort Barnwell Station and wind data from Cherry Point Marine Base

Weather Station (CP) and Cape Hatteras Weather Station (HT) was obtained from the

National Weather Service, NOAA. Asterisks indicate significant p-values.

Table 2. Three-way ANOVA comparisons of sea nettle count by site, month and year.

Asterisks indicate significant p-values.

Table 3. North shoreline, Pearson correlations with sea nettle count (log+1), in situ and

wind variables measured from Cherry Point Marine Base Weather Station (CP) and

Cape Hatteras weather station (HT); N = 137.

Table 4. South shoreline, Pearson correlations with sea nettle count (log+1), in situ and

wind variables measured from Cherry Point Marine Weather Station (CP) and Cape

Hatteras weather station (HT); N = 139.

Table 5. North shoreline, Pearson correlations with sea nettle count (log+1) and wind

events 3 to 8 ms-1 (N = 7) measured from Cherry Point Marine Base Weather Station

(CP)

49

Table 6. South shoreline, Pearson correlations with sea nettle count (log+1) and wind

events 3 to 8 ms-1 (N = 7) measured from Cherry Point Marine Base Weather Station

(CP)

Table 7. South shoreline, Pearson correlations with sea nettle count (log+1) and wind

events 2 to 7 ms-1 (N = 8) measured from Cape Hatteras Weather Station (HT)

Figure 1. (A) The Albermarle-Pamlico estuarine system (APES), which includes

Albermarle Sound, Pamlico River Estuary (PRE) and Neuse River Estuary (NRE). The

circles indicate the location of the North Carolina Division of Marine Fisheries oyster

sanctuaries. The Cape Hatteras Weather Station (HT) and Cherry Point Marine Base

Weather Station (CP) were the weather stations used to collect wind data for this study.

(B) Research was conducted in the Neuse River Estuary (NRE) at the 6 starred

locations: Camp Seafarer (CSF), Camp Sea Gull (CSG), Oriental Town (OM), Flanners

Beach (FB), Pine Cliff (PC), and Siddie Fields (SF). The squares within the NRE are the

approximate locations of the current nodes of the RENCI ADCIRC forecast model data

for 2011 (site nearest FB only) and 2012 (all gauges). The circles on this Figure indicate

the oyster sanctuaries in closest proximity to the NRE.

Figure 2. Box plots of stream flow and salinity in 2011 and 2012. T-test p-values are

displayed to compare differences in 2011 and 2012 mean stream flow (N = 231) and

salinity (N = 196), accordingly. Open circles indicate outliers > 1.5 times of upper and

lower quartile and horizontal bars within the box plots indicate the count median. The

50

upper box above the median extends to the third quartile and the lower box extends to

the first quartile. The bars extend to the largest and smallest values, respectively.

Figure 3. Box plots of wind speed and direction measured from Cherry Point Marine

Base (N = 170) and Cape Hatteras (N = 177) weather stations for 2011 and 2012. T-test

p-values are displayed to compare differences in 2011 and 2012 mean wind speed and

direction, accordingly. Horizontal bars within boxplots indicate sample median. The


the first quartile. The bars extend to the largest and smallest values, respectively.

Figure 4. The total amount of C. quinquecirrha counted with standard deviation error

bars for field seasons 2011 (N = 150) and 2012 (N = 149).

Figure 5. Box plots showing the range of sea nettle counts (Log +1) at the research

sites Camp Seafarer (CSF), Camp Sea Gull (CSG), Oriental Town (OM), Flanners

Beach (FB), Pine Cliff (PC) and Siddie Fields (SF) in 2011 and 2012. Open circles

indicate outliers > 1.5 times of upper quartile and horizontal bars within the box plots

indicate the count median. The upper box above the median extends to the third quartile

and the lower box extends to the first quartile. The bars extend to the largest and

smallest values, respectively. Data was standardized to ~100m observation length per

site.

51

Figure 6. Daily wind speeds and directions in the 2011 and 2012 field seasons at Cherry

Point Marine Base Weather Station. Inverted triangles indicate wind events, where

winds increased in speed from 3 to 8 ms-1.

Figure 7. Daily wind speeds and directions in the 2011 and 2012 field seasons at Cape

Hatteras Weather Station. Inverted triangles indicate wind events, where winds

increased in speed from 2 to 7 ms-1.

Figure 8. (A) Sea nettle abundance (m2) with standard deviation error bars compared to

wind directions observed at North shoreline sites Camp Seafarer (CSF), Camp Sea Gull

(CSG) and Oriental Town (OM) with wind events changed from 3 to 8 ms-1 occurring

five days prior to observations in July 2011, May 2012 and June 2012. (B) Sea nettle

abundance (m2) with standard deviation error bars compared to wind directions

observed at South shoreline sites Flanners Beach (FB), Pine Cliff (PC) and Siddie

Fields (SF) with wind events from 3 to 8 ms-1 occurring one day prior to observations in

July 2011, May 2012 and June 2012.

Figure 9. Conceptual diagram of wind-driven circulation circulation and NRE sea nettle

observations. Wide arrows indicate wind direction from the SW (A, C, D) and NE (B, E)

measured from Cherry Point Marine Base Weather Station (CP) or Cape Hatteras (HT)

and narrow arrows show water current direction based on work done by Luettich et al.

(2000), Reynolds-Fleming and Luettich (2004), the RENCI NC-CERA ADCIRC forecast

model data from 2012, and water depth observed in this study. Water depth (the

52

solid/curved line) at the coastline sites on the South shoreline (S), Flanners Beach (FB),

Pine Cliff (PC) and Siddie Fields (SF) and pier sites Camp seafarer (CSF), Camp Sea

Gull (CSG) and Oriental Town (OM) on the North shoreline (N) is influenced by wind-

driven circulation circulation in the NRE and may also affect sea nettle distribution and

abundance. Circles on each NRE map indicate the location of the two nearest oyster

sanctuaries. Shorelines, Pearson correlation coefficients and p-values are displayed on

the top of each figure and for figures D and E, above each abundance graph for

reference. The value and bar in the center of each abundance bar plot are the averages

of sea nettles counted at each site for both study years.

53

Table 1.

T-test comparison of abiotic variables by year variables year min max mean median t stat p-value Depth (m) 2011 0.3 2.7 1.3 1.0 -1.5 0.1 2012 0.3 3.1 1.4 1.2 Secchi (m) 2011 0.3 1.4 0.7 0.7 -1.6 0.1 2012 0.3 1.6 0.8 0.8 Temperature (ºC) 2011 22.2 38.0 29.5 19.2 7.0 < 0.001* 2012 21.3 33.9 27.5 28.9 Salinity 2011 8.8 30.7 18.9 19.2 13.7 < 0.001* 2012 1.0 23.7 13.1 13.6 Stream flow (m3s-1) 2011 425 2130 850 722 -12.1 < 0.001* 2012 559 6300 2469 2100 CP Wind speed (ms-1) 2011 1.0 8.0 3.9 4.0 3.6 0.0* 2012 1.0 7.0 3.2 3.0 CP Wind direction (North = 0°) 2011 48.0 246.0 161.9 180.0 1.7 0.1 2012 29.0 285.0 147.3 148.0 HT Wind speed (ms-1) 2011 1.7 6.4 3.8 3.6 -0.3 0.7 2012 2.0 7.0 3.8 4.0 HT Wind direction (North = 0°) 2011 46.3 258.3 176.6 194.2 0.8 0.5 2012 25 267.0 169.8 194.0

54

Table 2.

Comparison of sea nettle count by site, month and year ANOVA DF Sum Sq Mean Sq F-value p-value Site 5 21.6 4.3 23.1 < 0.001* Month 2 46.0 23.0 122.9 < 0.001* Year 1 0.34 0.3 1.8 0.18 Site * Month 10 19.8 2.0 10.6 < 0.001* Site * Year 5 3.7 0.7 4.0 0.00* Month * Year 2 7.8 0.9 4.8 0.00* Site * Month * Year 10 5.89 0.6 3.1 0.00*

55

Table 3. North shoreline, Pearson correlations with in situ and wind variables variables (log) R p-value stream flow (m3s-1) -0.20 0.01 CP 1 day prior wind speed (ms-1) 0.35 < 0.001 CP 2 day prior wind speed (ms-1) 0.28 0.00 HT 1 day prior wind speed (ms-1) 0.18 0.03 HT 2 day prior wind speed (ms-1) 0.25 0.00 HT 2 day prior wind direction (North = 0°) 0.19 0.02

56

Table 4. South shoreline, Pearson correlations with in situ and wind variables variables (log) R p-value depth (m) -0.19 0.02 secchi (m) -0.19 0.02 Salinity 0.18 0.04 CP 1 day prior wind speed (ms-1) 0.21 0.01 CP 1 day prior wind direction (North = 0°) 0.24 0.00 HT 1 day prior wind direction (North = 0°) 0.28 0.00 HT 2 day prior wind speed (ms-1) 0.22 0.00 HT 2 day prior wind direction (North = 0°) 0.20 0.02

57

Table 5. North shoreline, Pearson correlations with wind events 3 to 8 ms-1 (N = 7)variables (log) R p-value CP 5 days prior wind speed (m s-1) 0.41 0.03 CP 5 days prior wind direction (North = 0°) 0.53 0.00

58

Table 6. South shoreline, Pearson correlations with wind events 3 to 8 ms-1 (N = 7) variables (log) R p-value CP 1 day prior wind direction (North = 0°) 0.70 4.07 e-05 CP 3 days prior wind speed (m s-1) 0.57 0.00 CP 4 days prior wind speed (m s-1) 0.46 0.01 CP 6 days prior wind speed (m s-1) 0.53 0.00

59

Table 7.

South shoreline, Pearson correlations with wind events 3 to 8 ms-1 (N = 8) variables (log) R p-value HT 2 days prior wind direction (North = 0°) -0.54 0.01 HT 7 days prior wind speed (m s-1) 0.49 0.02

60

Figure 1.

61

Figure 2.

62

Figure 3.

63

Figure 4.

64

Figure 5.

65

Figure 6.

66

Figure 7.

67

Figure 8.

A

B

68

Figure 9.

69

70

CHAPTER 3

An assessment of the potential of sea nettle, Chrysaora quinquecirrha, sexual

reproduction in the Neuse River Estuary

72

ABSTRACT

This chapter focused on determining if sexual reproduction of the sea nettle,

Chrysaora quinquecirrha, was occuring the Neuse River Estuary (NRE). Many species

of jellyfish aggregate for sexual reproduction thereby potentially increasing the

encounter rates with humans. One-hundred jellyfish were randomly sampled from May-

July 2011 at six recreation sites. Histology was used to determine the sex ratio, the

presence/absence of mature gonads, and the presence/absence of brood planulae. The

jellyfish sex ratio was found to not be significantly different from 1:1. Based on egg

diameter, no females were sexually mature and no females had planulae present in

their gastric cavities. Of the 48 males, five showed rupture sperm follicles and all of

these males were sampled early in the season. There was no relationship between bell

and egg diameter, suggesting that female size was not related to egg maturity.

However, a negative linear relationship between bell diameter and time indicated a

gradual decrease in organism size over time. These results suggest 1) sexual

reproduction may occur very early in the season and sampling did not capture this event

or 2) sexual reproduction is not occurring and the primary source of medusae during the

year comes from the strobilation of polyps. These findings suggest that NRE C.

quinquecirrha egg maturity should be further investigated as these jellyfish may possess

smaller sized, mature eggs. Research on other life history stages (polyp, ephyrae, etc.)

should be conducted to gather more information on the longevity of C. quinquecirrha in

the NRE and greater APES. Further life history research will indicate which stage of

jellyfish, sexual or asexual, is more important to manage.

73

INTRODUCTION

This chapter evaluated the potential of sexual reproduction of Chrysaora

quinquecirrha as a potential factor related to jellyfish-human interactions in the Neuse

River Estuary (NRE). The annual occurrences of C. quinquecirrha may have negative

effects on NRE recreation and subsequently tourism revenue (Figure 1), which is

economically important to residents and NRE stakeholders. Estuaries can be highly

conducive to jellyfish populations by providing placid areas for adult jellyfish or medusae

to aggregate for spawning (Omori and Nakano 2001). The NRE is primarily influenced

by wind-driven circulation (see Chapter 2) and because C. quinquecirrha are weak

swimmers with maximum swimming rates of 1.8 cm s-1 (Costello et al. 1998; Matanoski

et al. 2001) and on average ~ 5 - 10 cm s-1 currents velocities are generated by wind

(Luettich et al. 2000; Reynolds-Fleming and Luettich 2004), aggregation of large

numbers of C. quinquecirrha by wind-driven circulation may or may not indicate specific

areas where sexual reproduction occurs in the NRE. Moreover, since the NRE is a

highly used recreation area, awareness that wind-driven circulation may influence

where sexual reproduction is likely to occur could help with managing this nuisance

species

The life history of Chesapeake Bay C. quinquecirrha populations has been

thoroughly studied (Figure 1). Pioneer research by Littleford (1939), Truitt (1939), Cargo

and Schultz (1966), Calder (1972, 1974), and Cargo and Rabenold (1980) have

documented the development and behavior of the C. quinquecirrha polyp, strobila, and

ephyrae in both laboratory and field conditions. In these studies, the asexual

reproduction phase of the life cycle is well described, including the perennial nature of

the polyp phase (Cargo and Schultz 1966; Truitt 1939) and how the polyp can

74

reproduce asexually through transverse fission (strobilation), pedal laceration, and/or

budding (Cargo and Rabenold 1980; Littleford 1939). In contrast, sexual reproduction of

the ephemeral medusa phase of the C. quinquecirrha life cycle has seldom been

described due to the difficult nature of documenting a spawning/fertilization event in situ,

which has been problematic with most jellyfish species (Arai 1997). Therefore a

traditional method to assess if C. quinquecirrha populations are reproducing sexually is

polyp-substrate studies whereby planulae settle on a substratum and undergo

metamorphosis into polyps (Breitburg and Fulford 2006; Cargo and Schultz 1966).

The best attempt to assess sexual reproduction in C. quinquecirrha was

performed by Littleford (1939). His research established a visual means of identifying

mature female and male C. quinquecirrha based on gonad color, how to determine

gonad maturity in female C. quinquecirrha, and presented a time frame from egg

fertilization to planula development within female C. quinquecirrha gastric cavities.

Spawning/fertilization occurred in the evening, from 2000-2100, and planula stages

were found from 1000 onward the next morning. Upon fertilization, development into

planula occurred instantly or started six or seven hours later. Planulae remained in the

gastric cavities no more than 24 hours after fertilization (Littleford 1939). Littleford

(1939) also suggested that fertilization and the development of planulae is more likely to

occur in the gastric cavities of female C. quinquecirrha than the water column (Figure

1B). However, a study on planulae behavior revealed that fertilization and development

outside female also occurs (Cargo 1979) (Figure 1A). To date, Littleford’s egg

fertilization and diameter maturity experiments have not been revisited and C.

75

quinquecirrha literature from the past ~ 35 years seldom report sex ratios of medusae,

gonad maturity, and if planulae are present in gastric cavities.

The primary objective of this chapter was to assess the potential of NRE C.

quinquecirrha sexual reproduction by using Littleford’s (1939) observations and sexual

reproduction characteristics reported from other jellyfish species. Chrysaora

quinquecirrha are gonochoristic and 1:1 sex ratios have been reported in other jellyfish

species known to sexually reproduce (Arai 1997; Pitt and Kingsford 2000; Rosa et al.

2012); therefore, I hypothesized that the proportion of female and male sea nettles

would not differ. Littleford’s (1939) experiments and observations showed that egg

diameters of 0.07 to 0.19 mm, with an average of 0.15 mm, could be successfully

fertilized and were classified as sexually mature (Littleford 1939). Based on these

observations, I evaluated gonad maturity in female NRE C. quinquecirrha and tested the

hypothesis that the majority of eggs found would be greater than 0.07 mm and therefore

sexually mature.

Jellyfish spermatogenesis occurs in follicles and spawning occurs when sperm

follicles rupture allowing mature sperm to enter the gastric cavity and dispensed orally

(Arai 1997). In addition to ruptured sperm follicles, an indication that sperm was

released into the water column can be determined by observations of “spent gonads,”

where instead of sperm the presence of amoeboid cells and free spaces in the lumen

are observed (Schiariti et al. 2012). To evaluate gonad maturity in male C.

quinquecirrha, I hypothesized that sexually mature males would have ruptured sperm

follicles or show signs of spent sperm follicles. It should be noted that spent gonad

observations have also been attempted on female jellyfish but due problems with

76

histology procedures, artifacts from staining slides make it difficult to discern spent

female gonads (Schiariti et al. 2012). Thus, I did not attempt to observe spent gonads in

female C. quinquecirrha. Internal fertilization of planulae occurs when sperm is taken up

from the water column into the gastric cavity by the female’s oral arms similar to feeding

(Arai 1997). If mature eggs are released from the female gonad at this time,

development of planulae occurs in the gastric cavity and the planulae will swim out of

the mouth of the females when fully developed (Littleford 1939). I hypothesized that

brooded planulae would be observed in the gastric cavity of females indicating that

sexual reproduction had occurred.

The size of jellyfish, which is typically measured by bell diameter, may influence

sexual reproduction and spawning commencement (Arai 1997) because jellyfish

maturation may be stopped and started as the medusa starves or eats and this change

in size is reflected in bell and egg diameter (Arai 1997). Experiments with the moon

jellyfish Aurelia sp. documented that when the jellyfish shrink or “degrow” the female

gonad regresses along with egg maturity. Spermatogenesis, however, continues to

proceed regardless of gonad regression (Hamner and Jenssen 1974). In recent studies

where jellyfish were surveyed in situ, female egg diameter, fecundity, and bell diameter

were positively linear (Saucedo et al. 2012) or unrelated to the size of jellyfish

(Toyokawa et al. 2010). To date, no comparisons with bell and egg diameter have been

made with C. quinquecirrha. I hypothesized that bell diameter would not be correlated to

egg diameter in female C. quinquecirrha. If bell diameter and egg diameter are

correlated, it would be possible to determine sexual maturity of females from size alone,

to the exclusion of histology. In addition, since Littleford (1939) observed mature

77

females to have grayish-yellow brown gonads and males with bright pink colored

gonads, I also explored the possibility of identifying sex of C. quinquecirrha in situ by

taking observations of gonad color.

The populations of C. quinquecirrha in the NRE are influenced by wind dynamics

(see Chapter 2). In this chapter, I evaluated if these wind-driven aggregations may be

related to the potential of sexual reproduction of NRE C. quinquecirrha thereby

contributing to the annual occurrences of jellyfish-human interactions in the NRE. In

response to the ecological and societal implications of large numbers of jellyfish in

coastal environments worldwide, sexual reproduction capabilities in jellyfish species

have steadily gained attention (Saucedo et al. 2012; Schiariti et al. 2012; Toyokawa et

al. 2010). Research from this chapter was the first to assess sexual reproductive

characteristics in NRE C. quinquecirrha. I anticipate that these results will contribute to

jellyfish reproduction studies and on-going jellyfish research in North Carolina.

MATERIALS AND METHODS

Gonad collection and histology

A total of 100 C. quinquecirrha were randomly collected in 2011 on 19 May, 3

June, 10 June, 18 June, 8 June, 5 July, 12 July, and 19 July by dip netting (Figure 2B).

Basic morphological data including bell diameter, bell color, oral arm color and visual

observations of the sex of medusa were recorded for each collected C. quinquecirrha.

Bell and oral arm colors were also documented because there are different color

varieties of C. quinquecirrha in the NRE, similarly to Chesapeake Bay C. quinquecirrha

(Littleford 1939). The entire gastric cavity (the mouth, stomach, and four gastric

78

pouches) excluding oral arms was dissected out of each jellyfish and preserved in 5%

formalin in filtered estuarine water. All gonad samples were collected twice a week from

the hours of 0800 and 1700; therefore, I presumed all female gonad observations would

have an assortment of eggs, mature eggs, and/or planulae based on Littleford’s (1939)

fertilization and planulae development timeline. Since each gastric pouch contains a

gonad with the potential of harvesting mature eggs, planulae or sperm, all four gonads

were placed in a single histology cassette and a single slide was created per jellyfish.

Additionally, C. quinquecirrha sperm is located in one of the four gastric pouches

(Littleford 1939) so analyzing all four gonad pouches assured that I could properly

identify if the jellyfish was male or female. Tissue samples of fixed gonads were washed

in an ethanol series, embedded in paraffin and 5 μm slices were placed on each

histology slide. Histology slides where stained with a Harris Hematoxylin and Eosin

stain (Edna and Prophet 1992) and all histological procedures were performed at the

East Carolina University Histology Core Facility at the Brody Medical School, Greenville,

North Carolina.

Microscopy and CellSens image analysis

All slides were analyzed with an Olympus BX41 light microscope at 10x to 40x

magnification. Gonad pictures were taken and analyzed with CellSens image analysis

software. To determine how many egg diameters were needed to account for sample

variation, the diameters of all eggs were measured for six jellyfish. The number of eggs

per jellyfish ranged from 106 to 4,137, and a standard deviation analysis of the

measured egg diameters showed that fluctuations in measurements plateaued when

79

100 egg diameters per slide were measured. Therefore, for the remaining slides with

eggs, approximately 100 randomly selected egg diameters were measured. Eggs were

identified by a well-defined nucleus, circular shape and egg diameters were measured

at the widest part of each egg (Figure 3A). Fertilized eggs were identified by

multicellular composition and larger diameters. If planulae were present, the number of

planulae was recorded within each of the four gastric pouches. Identification of planulae

was based on the oval shaped characteristic of the larvae and multicellular composition.

If sperm follicles were present (Figure 3B), I searched the entire gonad for ruptures in

cell membranes or spent sperm follicles as described by Schiariti et al. (2012) (Figure

4). If ruptured sperm follicles or spent sperm follicles were present, these individuals

were classified as mature males.

Statistical analysis

To meet the requirements for parametric tests, all measured egg diameters

belonging to each female jellyfish were Log transformed. To evaluate sex ratio, a chi-

square test was performed on all jellyfish collected and distributed by site and

throughout the field season. I used a one-way t-test to evaluate if the egg diameters

were > 0.07 mm. Bell diameters were also logged transformed and an ANOVA was

used to compare egg diameters by site, bell diameters by site, and bell diameters by

maturity. Bartlett tests of variance were performed to evaluate the validity of each

ANOVA and if significant, Tukey post hoc tests were used to compare means. Linear

regression analyses were used to compare female bell and egg diameters and bell

80

diameters of females and males throughout the field season. All statistical analyses

were performed with R version 2.14.2 (2012-02-29).

RESULTS

Sex ratios and microscopy observations

I found a sex ratio of 13:12 with 52% females and 48% males, which is not

significantly different that a 50:50 ratio (χ2 = 0.16, DF = 1, p-value = 0.69). The

frequency of females and males found at each site was relatively uniform except for FB

(Figure 1B) where there were ~65% females and ~35% males (χ2 = 9, DF = 1, p-value =

0.00) (Figure 5). The frequency of females and males found throughout the field season

also had a somewhat uniform distribution, however more males were observed on 19

May (χ2 = 36, DF = 1, p-value = 2.0 e-09), and 17 July (χ2 = 16, DF = 1, p-value = 6.3 e-

05) and more females were observed on 1 June (χ2 = 16, DF = 1, p-value = 6.3 e-05)

and 12 July (χ2 = 8.6, DF = 1, p-value = 0.00) (Figure 6).

Based on the egg diameter maturity of > 0.07 mm (Littleford 1939), none of the

females sampled in our study were sexually mature (Figure 7). The ANOVA of mean

egg diameters by site was not significant (Table 1), but the range of egg diameters was

higher at the South sites (FB, PC and SF) and at CSG (Figure 1B, Figure 8). Sites CSF

and FB had the largest range in egg diameter (Figure 8). No planulae were observed in

the gastric cavities. Of the males, five showed ruptured sperm follicles and sperm

entering the gastric cavity of the male jellyfish (Figure 4). Males with ruptured sperm

follicles were found early in the field season and at sites OM and PC (Figure 1B) and I

81

did not observe spent sperm follicles any of the male samples. Unlike Littleford’s (1939)

observations, I found sperm follicles in all four gonads for each male C. quinquecirrha.

Bell diameter comparisons

The range of bell diameters and mean bell diameter for females, immature males

and mature males varied at each site (Figure 9A, Figure 9B and Figure 9D, Table 1).

Immature females and males had a similar range in bell diameters (cm) and medians,

which seem to be smaller than the mature males (Figure 9C). The range of mature

males was between 7 - 12 cm at OM and ~ 8 - 10 cm at PC (Figure 9D), but the ANOVA

of maturities was insignificant. Therefore, it is difficult to compare if the mean bell

diameters of immature and mature males are similar or dissimilar (Table 1). The

ANOVAs and Bartlett tests of variance were significant for the female and immature

male bell diameters by site comparisons (Table 1). The Tukey post hoc test of mean

bell diameters of females were significantly different between OM, north, and most of

the south sites (CSF, p-value = 0.01; CSG, p-value = 0.00; FB, p-value = 0.00; SF, p-

value = 0.00). Mean bell diameters of immature males were also significantly different

between OM and CSG (p-value = 0.00), and all south sites (FB, p-value = 0.00; PC, p-

value = 0.02; SF, p-value = 0.01) (Figure 9A, 9B).

Relationship between egg and bell diameters

There was no linear relationship between egg and bell diameter (r2 = 0.01, p-

value = 0.5, Figure 10); however, there was a negative linear relationship between

female bell diameter and time throughout the field season and a positive linear

82

relationship between egg diameter and time throughout the field season (Figure 11).

There was also a negative linear relationship between male bell diameter and time

throughout the field season (Figure 12).

Histology versus in situ gonad observations

Sex determined by histology was substantially different than in situ gonad

observations to classify sex. Despite the differences in bell and oral arm color, females

were more accurately identified than males. There was no bell and oral arm color

combinations that were associated with the sex of jellyfish (Table 2).

DISCUSSION

I found a 1:1 sex ratio (Figure 5 and Figure 6) and of the male C. quinquecirrha

surveyed ~10% showed ruptured sperm follicles, which indicated that these males were

sexually mature. These individuals were collected on 19 May and 10 June, when egg

diameters were measured at the 0.035 to 0.040 mm (Figure 11, Figure 12), which was

the most frequently measured egg diameter in my study (Figure 7). Mature males were

also collected on 3 June at OM and PC when egg diameters were smaller, 0.010 to

0.030 mm (Figure 11, Figure 12). The ANOVA of mean egg diameters by site was not

significant (Table 1) but the range of egg diameters was higher at the South sites (FB,

PC and SF) and at CSG (Figure 1B, Figure 8). Sites CSF and FB had the largest range

in egg diameter (Figure 8). None of the females sampled were sexually mature based

on Littleford’s (1939) egg diameter index but it is possible that mature egg diameters

may be smaller for NRE C. quinquecirrha populations. Recent histology studies

83

evaluating reproductive capabilities in Rhizostomae jellyfish known as broadcast

spawners have used egg yolk content and egg diameters to evaluate egg maturity in the

giant jellyfish Nemopilema nomurai [Kinoshinouye, 1922] (Iguchi et al. 2010; Toyokawa

et al. 2010) and Lychnorhiza lucerna [Hackel, 1880] (Schiariti et al. 2012). For N.

nomurai, histology and fertilization experiments have indicated that smaller egg

diameters could be successfully fertilized under favorable conditions with adequate yolk

content (Kawahara et al. 2006; Ohtsu et al. 2007; Toyokawa et al. 2010). If this is the

case with C. quinquecirrha, and mature eggs were present within the females surveyed,

then the presence of mature males could indicate that commencement and the potential

of spawning occurred early in the field season potentially at PC and OM where the

mature males were collected.

Another way to directly document reoccurring reproduction events is the

observation of fertilized eggs, embryos, and planulae inside female gastric cavities of

brooder jellyfish species (Brewer 1989; Schiariti et al. 2012). Since I did not find

brooded fertilized eggs and/or planulae, there was no evidence that egg fertilization had

occurred in the gastric cavity among the females I sampled; however, it is also possible

that fertilization and planulae development occurred in the water column, similar to other

broadcast spawning jellyfish species (Figure 2A) and noted by Cargo (1979) for C.

quinquecirrha. Furthermore, if egg production rates are similar to Chesapeake Bay C.

quinquecirrha (up to 40,000 eggs daily (Purcell, unpublished data)), the overturn of

immature to mature eggs could be on the order of hours and mature eggs may have

been expelled prior to capture of the female. Internal planulae development (Figure 2B)

and departure may have also been missed. On the other hand, since my sampling was

84

conducted bi-weekly and at times that coincided with Littleford’s (1939) observations of

fertilization and planulae development, the possibility that sexual reproduction did not

occur is also possible, assuming NRE sea nettles adhere to similar fertilization and

development time frames as documented in Chesapeake Bay.

I found that bell and egg diameters were not linearly related (Figure 10) but,

similar to Toyokawa et al. (2010), I observed an increase in egg diameter as the season

progressed (Figure 11). Although no egg diameters were measured, unpublished data

have suggested that C. quinquecirrha in Chesapeake Bay are sexually mature at bell

diameters approximately 2 cm (Purcell et al. 1999). Based on this finding, I could

classify all C. quinquecirrha sampled in our study as sexually mature and this could

explain why there was no relationship between bell and egg diameters (r2 = 0.01, p-

value = 0.5, Figure 10). Mean bell diameter of females and immature males was

significantly different among sea nettles collected from OM when compared to the other

sites (Figure 9A, 9B). Specifically, the mean bell diameters of these jellyfish appear to

be smaller than those collected elsewhere. The negative linear relationship between bell

diameters and time (Figure 11 and Figure 12) may reflect the seasonality of C.

quinquecirrha where medusa numbers decline in late summer (Decker et al. 2007) and

in conjunction with observed decreases in bell diameter (Purcell 1992).

My in situ observations of sex by gonad color proved to be misleading (Table 2);

therefore, to determine sex in C. quinquecirrha, histology procedures should be used.

These results also concur with other jellyfish reproduction histology observations where

gonad color varied with sex in N. nomurai (Toyokawa et al. 2010) and L. lucerna

(Schiariti et al. 2012). However, it is also possible that because most of the C.

85

quinquecirrha surveyed were classified as sexually immature, the coloration of mature

gonads described by Littleford (1939) may have not been observed because the jellyfish

were, in fact, not sexually mature. Lastly, the different color varieties of bell and oral arm

color observed in my study showed no color combination associated with sex and these

observations coincide with Littleford’s (1939) conclusions of bell color, oral arm color,

and sex.

CONCLUSIONS

The use of histology proved to be an informative way to assess the potential of

sexual reproduction of C. quinquecirrha in the NRE. Although my results cannot

definitively support that the annual occurrences of C. quinquecirrha are related to

aggregation for the purposes of sexual reproduction, I can report that 1:1 sex ratios

were observed at all recreation sites throughout the 2011 season and mature males

were collected early in the field season at a North (OM) and South (PC) shoreline site

(Figure 1B, Figure 9D). These observations support the argument that sexual

reproduction could occur in the NRE and in areas that are used for recreation and/or

tourism, but with equal evidence that sexual reproduction did not occur in 2011 (i.e.,

immature females based on egg diameter and no brooded planulae). I suggest that egg

maturity be revisited before classifying what recreation sites or NRE shorelines are

more prone to C. quinquecirrha aggregations for sexual reproduction. Future studies

should include a variety of methodologies including histology, in vitro fertilization

experiments, and the use of ecological techniques. For example, settlement plate

studies similar to those conducted in Chesapeake Bay (Calder 1972; Cargo and Schultz

86

1967) and with other jellyfish species (Astorga et al. 2012; Holst and Jarms 2007;

Hoover and Purcell 2009) should be conducted to investigate the potential of planulae

settlement as a proxy for sexual reproduction events as well as substrate preference.

Studies have suggested that C. quinquecirrha planulae prefer to settle on natural

substrates, such as oyster shells (Breitburg and Fulford 2006; Purcell 2012) but other

studies have shown that artificial substratum may also be selected by planulae of other

species (Lo et al. 2008; Toyokawa et al. 2011). As discussed in chapter 2, if NRE C.

quinquecirrha planulae prefer to settle on oyster shells like Chesapeake Bay C.

quinquecirrha, the NC Division of Marine Fisheries oyster sanctuaries distributed

throughout APES (Figure 1A) could serve as adequate habitats. In addition, if C.

quinquecirrha planulae settled on artificial substrates, there are many docks and piers in

the NRE that could also serve as suitable substrate.

Ample habitat for settlement would support the stability of the asexual life history

stages (polyp and strobila) thereby increasing the likelihood of continued annual

occurrences of C. quinquecirrha in the NRE driven by the asexual reproduction

component of the bipartite lifecycle. For example, the polyp stage is highly resilient to

changes in low dissolved oxygen (Condon et al. 2001), will cyst when unfavorable

conditions are present, including changes in season and/or water temperature (Cargo

and Schultz 1966), and may survive for several years (Cargo and Schultz 1967).

Therefore, stable polyp populations that undergo metamorphosis into ephyrae each

spring (Calder 1974; Cargo and Rabenold 1980) are not dependent on sexual

reproduction making the possibility of NRE C. quinquecirrha annual medusae

occurrences low in genetic diversity, but harmful to humans nonetheless. Genetic

87

surveys of clonal generations and inference to reproductive mode of organisms

belonging to Phylum Cnidaria have been conducted with the Hydrozoa (hydromedusae)

(Meek et al. 2013) and Anthozoa (corals) (Jokiel et al. 2013) and similar techniques

could be used to assess clonal diversity in the Scyphozoa or jellyfish similar to C.

quinquecirrha.

The results reported in Chapter 2 and 3 seem to indicate that the physical driver

of wind-driven circulation is more influential to determining the extent and frequency of

jellyfish-human interactions in the NRE than sexual reproduction. Therefore, coastal

management of sea nettles in the NRE should focus on mitigation strategies as outlined

in Chapter 2 to minimize harmful jellyfish-human interactions with C. quinquecirrha,

including barrier nets (Schultz and Cargo 1969), public outreach, awareness and first

aid to treat stings as demonstrated in Australia (Gershwin et al. 2010) and Germany

(Baumann and Schernewski 2012) instead of biological control methods such as

removal of the medusae from the system.

88

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Table 1. Analysis of variance and Bartlett variance tests for egg diameters (mm) by site,

bell diameters (cm) by site and maturities (immature females, immature males, and

mature males). Asterisks indicate significant p-values.

Table 2. Bell and oral arm color comparisons with in situ gonad observations and sex of

females (F) and males (M) determined by histology.

Figure 1. (A) Conceptual diagram of the life cycle of C. quinquecirrha where spawning

of gametes, fertilization and development of planula occur in the water column. (B)

Conceptual diagram of the life cycle of C. quinquecirrha where sperm is taken up from

the water column into the female’s gastric cavity, where fertilization and development of

planula occurs.

Figure 2. (A) The Albermarle-Pamlico estuarine system (APES), which includes

Albermarle Sound, Pamlico River Estuary (PRE) and Neuse River Estuary (NRE). The

circles indicate the location of the North Carolina Division of Marine Fisheries oyster

sanctuaries. (B) Jellyfish were collected from the Neuse River Estuary (NRE) at the 6

starred locations: Camp Seafarer (CSF), Camp Sea Gull (CSG), Oriental Town (OM),

Flanners Beach (FB), Pine Cliff (PC), and Siddie Fields (SF) from Mid-May to July 2011.

The circles indicate the oyster sanctuaries in closest proximity to the NRE.

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Figure 3. Microscopy images of NRE C. quinquecirrha eggs (A) and sperm follicles (B).

Figure 4. Microscopy images of NRE C. quinquecirrha ruptured sperm follicles with

arrows indicating sperm entering the gastric cavity.

Figure 5. Observed sex ratio at each site; Camp Seafarer (CSF), Camp Sea Gull

(CSG), Oriental Town (OM), Flanners Beach (FB), Pine Cliff (PC) and SF (Siddie

Fields).

Figure 6. Sex ratio observed throughout 2011 season. Mature males were collected on

19 May (N = 1), 3 June (N = 3) and 10 June (N = 1).

Figure 7. Histogram of all 52 female C. quinquecirrha surveyed. Egg diameters

measured between 0.035 and 0.040 mm were the most frequently observed.

Figure 8. Box plot of female egg diameters (N = 52) observed at each site; Camp

Seafarer (CSF), Camp Sea Gull (CSG), Oriental Town (OM), Flanners Beach (FB), Pine

Cliff (PC) and SF (Siddie Fields). Open circles indicate outliers > 1.5 times of lower

quartile and horizontal bars within the box plots indicate the egg diameter median. The


the first quartile. The bars extend to the largest and smallest values, respectively

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Figure 9. Box plots of immature female (N = 52, A), immature male (N = 43, B) and

mature male (N = 5, D) bell diameters observed at each site; Camp Seafarer (CSF),

Camp Sea Gull (CSG), Oriental Town (OM), Flanners Beach (FB), Pine Cliff (PC) and

SF (Siddie Fields), throughout 2011. (D) Box plots of bell diameters of immature

females, immature males and mature males. Open circles indicate outliers > 1.5 times

of upper quartile and horizontal bars within the box plots indicate the bell diameter

median. The upper box above the median extends to the third quartile and the lower

box extends to the first quartile. The bars extend to the largest and smallest values,

respectively

Figure 10. Scatter plot of female egg (mm) and bell diameters (cm), r2 = 0.01, p-value =

0.5

Figure 11. Scatterplots of female bell (diamond) and egg (circle) diameters from May to

July 2011. Linear regression lines show a decrease in bell diameter through season (r2

= 0.3, p-value = 0.0, black dashed line) and an increase in egg diameter (r2 = 0.2, p-

value = 0.0, grey line).

Figure 12. Scatterplot of male bell diameters from May to July 2011. Arrows indicates

the five mature males surveyed in this study. Similar to female bell diameter, male bell

diameter decreased over the field season (r2 = 0.3, p-value = 0.00, black regression

line).

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Table 1.

ANOVA Analyses by site, egg (mm), and bell diameters (cm) by site DF Sum Sq Mean Sq F-value p-value Bartlett variance test Female egg diameters 5 0.00 0.00 1.3 0.3 none performed Female bell diameters 5 121.8 24.4 8.1 < 0.001* p-value = 0.6 Male bell diameters 5 76.6 15.3 4.8 0.00* p-value = 0.8 ANOVA Analyses by bell diameters and maturities by maturities DF Sum Sq Mean Sq F-value p-value Bartlett variance test All bell diameters 2 21.6 10.8 2.2 0.11 none performed

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Table 2.

Histology versus in situ gonad observations bell color / oral arm color

sex determination via histology clear / clear clear / red red / clear red / red

F M F M F M F M 20 16 24 22 3 4 5 6

# of accurate in situ sex determination clear / clear clear / red red / clear red / red 20 1 23 0 3 0 4 0

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Figure 1.

A

B

100

Figure 2.

101

Figure 3.

A

B

102

Figure 4.

103

Figure 5.

104

Figure 6.

105

Figure 7.

106

Figure 8.

107

Figure 9.

A B

D C

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Figure 10.

109

Figure 11.

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Figure 12.

CHAPTER 4

Predators, stingers and economic influencers: a cultural consensus analysis of public

perception and ecological knowledge of jellyfish

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ABSTRACT

This chapter evaluated the social drivers of jellyfish-human interactions in

Eastern North Carolina. Large numbers of jellyfish have created many ecological and

societal problems worldwide. Specifically, adverse effects on fisheries and tourism have

been observed when jellyfish-laden waters have interfered with fisheries operations or

harmed people that encounter stinging species. Jellyfish populations exist on majority of

U.S. coastlines and in some areas increases have been observed. Coupled with more

coastal development, it is likely that U.S. coastal economies and communities will be

affected by jellyfish. To successfully manage these jellyfish-human interactions,

quantitative data on the public perspective of jellyfish ecology and how jellyfish influence

society is needed. This chapter used “cultural consensus theory” (CCT) to compare

public perspective of jellyfish across four culturally distinct groups of people: fishers

(commercial and recreational), recreationists, coastal, and jellyfish researchers. When

cultural knowledge of jellyfish ecology was compared, jellyfish researchers had the

highest cultural competency but similar mean cultural competencies between coastal

and jellyfish researchers were found. Mean cultural competencies among fishers,

recreationists and coastal researchers were also similar. When shown food web

illustrations, jellyfish researchers placed jellyfish in higher trophic levels in comparison

to fishers, recreationists and coastal researchers who placed jellyfish at lower trophic

levels. This could indicate that there is confusion as to the roles that jellyfish play in food

webs. All groups agreed that fewer than five jellyfish will not affect people’s decisions to

engage in water activities but the presence of jellyfish does affect people’s decisions to

continue water activities, especially if stung by jellyfish. However, people will book

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return vacations to areas known to experience jellyfish, even if they are stung. This

chapter revealed that more education is needed about the role of jellyfish in food webs.

Also, the knowledge that a threshold of tolerance (i.e., fewer than five) exits regarding

jellyfish in coastal areas utilized for water activities could be used for tourism planning

through coping and mitigation strategies such as on-site first aid, education and barrier

nets for designated swimming areas.

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INTRODUCTION

Jellyfish (Phyla Cnidaria and Ctenophora) are intermediate to top trophic level

predators in coastal and estuarine food webs (Mills 1995; Purcell et al. 2007). In these

environments, jellyfish prey heavily upon different species of zooplankton, including fish

larvae and eggs (Arai 1988; Moller 1984; Purcell 1992; Purcell and Arai 2001; Purcell

and Sturdevant 2001) and other jellyfish (Baird and Ulanowicz 1989; Purcell and Cowan

1995; Purcell and Decker 2005). Jellyfish predation affects fish populations as jellyfish

are often in direction competition with zooplanktivorous fish for similar prey, also when

large amounts of prey are present jellyfish feed without apparent satiation (Deason and

Smayda 1982; Hay 2006; Kremer 1979). Therefore, increases in jellyfish abundance

can result in the rapid depletion of prey resources to the detriment of fish. Jellyfish also

feed upon fish eggs and larvae, directly impacting fish populations in their early life

history (Purcell et al. 1994). Jellyfish are consumed by roughly 124 fish species and 34

other animal species (Arai 1988; Pauly et al. 2009; Purcell and Arai 2001), such as the

endangered leatherback turtle Dermochelys coriacea (Eisenberg and Frazier 1983;

Houghton et al. 2006), seabirds (Harrison 1984), crustaceans (Pauly et al. 2009)

cephalopods (Heeger et al. 1992) and humans (Hsieh et al. 2001; Omori 1978; Omori

and Nakano 2001; Pitt 2010). However, jellyfish are not consumed by predators as

readily as fish. In consequence, carbon is accumulated in jellyfish biomass and trophic

transfer to other, higher trophic level organisms is substantially reduced. The result is an

altered food web with a larger fraction of biomass consisting of jellyfish (Brodeur et al.

2011; Condon et al. 2011).

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The importance of jellyfish in ecosystems is frequently overgeneralized,

misconstrued, and/or not well-known to the public (Condon et al. 2012; Purcell et al.

2007). When compared to other species, jellyfish are typically understudied (Hay 2006),

especially in estuaries where jellyfish are intermediate-top level predators (Condon and

Steinberg 2008). More research is needed to further understand how jellyfish affect

pelagic and benthic food webs, how the degradation and utilization of jellyfish biomass

influences large scale biogeochemical processes (Ducklow et al. 2009), and the positive

benefits that jellyfish predation in food webs could have on biodiversity (Condon et al.

2011). Scientists often exclude jellyfish in ecological studies due to difficulties in

sampling (Purcell 2009) and some researchers have labeled jellyfish as “nuisance

species” (Richardson et al. 2009).

Occurrences of high jellyfish abundances (sometimes called blooms) have

caused many socio-economic problems globally. For example, Japan and India have

experienced power outages due to jellyfish clogging cooling intake-valves of coastal

power plants (Matsueda 1969; Matsumura et al. 2005; Rajagopal et al. 1989; Yasuda

1988). Jellyfish have interfered with aquaculture operations by feeding on reared

animals in Asia, Australia/Indo Pacific, Europe and North America (see Purcell et al.

2007 for review). In addition, fisheries in North America (Graham et al. 2003), the Black

Sea (Darvishi et al. 2004; Daskalov 2002; Shiganova et al. 2003; Zaitsev 1992),

Namibia (Lynam et al. 2006) Japan (Uye 2011), China (Dong et al. 2010), Brazil

(Nagata et al. 2009), Argentina (Schiariti et al. 2008), and Peru (Quinones et al. 2012)

have also suffered from excessive amounts of jellyfish that clog and burst nets.

Overfishing also alters food webs that favor jellyfish abundance instead of commercial

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fish species (Boero et al. 2008; Pauly et al. 1998; Purcell 2012). Seasonal occurrences

of jellyfish can also be problematic to fisheries. For example, Chrysaora plocamia

[Lesson, 1832] abundances peak in the summer and can be caught as 30% by-catch in

5% of Peruvian anchovy Engraulis ringens [L. Jenyns, 1842] hauls. This equates to an

economic loss of over US$ 200,000 in about a month’s worth of time and fishery

factories have refused to process hauls if catches include more than 40% jellyfish by-

catch (Quinones et al. 2012). The weight of jellyfish by-catch has also increased the risk

of capsizing trawling vessels (Graham et al. 2003). Processing hauls with jellyfish

requires more labor (Kawahara et al. 2006b) and because jellyfish sting, fish catch

mortality has increased (Bamstedt et al. 1998) and fish handlers who inevitably touch

the jellyfish while working are stung (Kawahara et al. 2006a).

Jellyfish are also a great concern to coastal areas that rely on tourism for

revenue (Purcell et al. 2007; Richardson et al. 2009). Periodic beach closures and injury

(including death) to recreationists or people that spend a long time in the water

swimming, wading, surfing, etc., have occurred globally in France, Spain, Thailand, and

Australia (Fenner et al. 2010; Fenner and Williamson 1996; Gershwin et al. 2010;

Pages 2001). Jellyfish stings in tourism areas have been an on-going coastal

management problem in Australia; however, public awareness and mitigation strategies

have been adopted to alleviate detrimental tourism effects (Gershwin et al. 2010).

Public education and awareness, including information posted on signs and pamphlets

about jellyfish given to beach-goers, have also helped people cope with large numbers

of jellyfish on German beaches (Baumann and Schernewski 2012). In Chesapeake Bay,

a monitoring system is in place that predicts the likelihood encountering a sea nettle

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(Decker et al. 2007). In other areas without extensive jellyfish research, public

monitoring of high numbers of stinging species has been conducted to reduce harmful

jellyfish-human interactions. Examples of these observations include the harmful box

jellyfish species in Hawai‘i (Thomas et al. 2001), historically referred to as Carybdea

alata [Reynaud, 1830] but currently regarded as Alatina moseri [Mayer, 1906] (Bentlage

et al. 2010; Gershwin 2005) and more recently the sea nettle C. quinquecirrha in

Barnegat Bay, New Jersey (Vasslides, pers comm.). Although loss of tourism and/or

recreation revenue due to jellyfish has not been recorded in these coastal areas or other

areas with high jellyfish abundance, researchers have proposed that jellyfish-infested

beaches will affect tourism (Purcell et al. 2007). Moreover, current media perspectives

on jellyfish populations in the environment are often negative (Condon et al. 2012) and it

has been suggested that as human populations and coastal recreation increase,

jellyfish stings will continue to be problematic (Macrokanis et al. 2004).

To date, jellyfish populations have increased in certain regions of the world

(Condon et al. 2012, Condon et al. 2013) and along the majority of U.S. coastlines,

including Hawai‘i and Alaska, jellyfish populations have shown varying degrees of

increase and certainty (Brotz et al. 2012). For example, an increase in jellyfish

populations was documented with high certainty along the U.S. Northeast coast and

Hawai‘i versus low certainty of increased jellyfish populations along the U.S. West coast

and Gulf of Mexico (Brotz et al. 2012).There are many hypotheses associated with

human activities as drivers for increasing jellyfish populations, including cultural

eutrophication, habitat modification, transportation of ballast water, aquaculture

practices and overfishing (See Purcell 2012 for review). Additionally, jellyfish undergo

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natural bloom and burst cycles of abundance, which can explain why large numbers of

jellyfish are observed in periodic and seemingly aberrant fluctuations (Boero et al.

2008). Although these jellyfish ‘blooms’ have been described as ecological enigmas

these occurrences are not ephemeral; instead, the ‘blooming’ nature of jellyfish species

is a product of the bipartite lifecycle (Müller and Leitz 2001). Therefore, in order to

accurately report jellyfish population dynamics instead of blooming events, more long

term data sets of jellyfish abundance is needed (Purcell et al. 2007).

Excluding the Great Lakes, about 30% of the total U.S. population resides on the

coast (Crowell et al. 2007). Moreover, the coastal population of the United States has

increased from 275 to 400 people per square kilometer from 1960 to 1990; by 2025, it is

predicted that nearly 75% of U.S. citizens will live in a coastal county or within 150km

from the coast (Hinrichsen 1995). This increase in coastal communities and the

existence of jellyfish populations along almost all U.S. coastlines equates to a

considerable likelihood of jellyfish-human interactions. People-wildlife interactions and

the rate of change in people’s beliefs and attitudes about human-environment relations

creates challenges in wildlife management (Decker and Enck 1996) and the efficient

sustainability of environmental resources (Cater 1995). Therefore, to circumvent

negative societal repercussions, such as loss of tourism revenue due to jellyfish

encounters and/or not utilizing coastal areas due to jellyfish, determining the current

state of jellyfish-human interactions in coastal communities affected by jellyfish is

needed.

I chose to investigate jellyfish-human interactions in North Carolina by analyzing

public perspective or cultural knowledge of jellyfish ecology and how jellyfish influenced

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people’s decisions to engage in water activities with cultural consensus theory (CCT).

CCT is a theory that forms the foundation for cultural consensus analysis, which is a

method used in anthropology that allows researchers to analyze qualitative data with

quantitative data analyses to estimate cultural beliefs and to report an individual’s

knowledge of those beliefs, known as cultural competency (Weller 2007). The overall

objective of this chapter was to understand public perspectives of jellyfish. Specifically, I

was interested in how cultural perspectives or knowledge of jellyfish would differ

between four culturally distinct groups; fishers (recreational and commercial),

recreationists, coastal researchers, and jellyfish researchers. These groups were

selected for this study because of the high likelihood of interacting with jellyfish while

engaging in water activities, including research. I evaluated jellyfish-human interactions

of these groups by distributing a jellyfish survey that tested two hypotheses related to

jellyfish ecology and how jellyfish influence society with an emphasis on people’s choice

to engage in water activities. When compared to other ecosystem organisms, jellyfish

are often understudied despite being intermediate to top level predators (Figure 1),

therefore, it is likely that ecological literacy of jellyfish is misinterpreted by researchers

other than jellyfish researchers. Moreover, since outreach education and associated

social media about scientific explorations stems from research (McKenna and Main

2013), misinterpretation of jellyfish ecological literacy is possible and may reverberate to

the public interface. To test this phenomenon, I hypothesized that the cultural

perspectives of jellyfish ecology would not differ among fishers, recreationists, coastal

researchers, and jellyfish researchers.

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I wanted to gather insights on how people are influenced by jellyfish, including

cultural perspectives of jellyfish stings, myths, and how jellyfish influence people’s

choices to engage in water activities. I chose to include common myths about jellyfish in

North Carolina because residents reported to us that jellyfish of particular colors will

“sting differently.” I also included common myths about jellyfish such as urination as a

treatment for jellyfish stings, where stings are felt on a person’s body and how jellyfish

will “appear” without warning. Jellyfish have influenced coastal economies that rely on

tourism revenue; therefore, I also wanted to explore how jellyfish-human interactions

affect water recreation, hobbies, and vacations to areas prone to jellyfish occurrences.

Each group was presumed to have different experiences and beliefs of jellyfish, thus,

perspectives of jellyfish-human interactions would also be variable among the groups.

For example, jellyfish researchers may perceive that large amounts of jellyfish are

tolerable in coastal waters because they study jellyfish versus recreationists may

perceive that large amounts are not tolerable because jellyfish inhabit desirable surfing

areas. Moreover, fishermen and coastal researchers may perceive jellyfish as problems

because of gear interferences and/or damage. To account for this variation in

perspectives, I hypothesized that the Cultural perspectives of jellyfish in society would

be similar among the cultural groups.

Data on the public’s perspective of unknown environmental-societal questions,

problems and/or concerns has been used to help manage environmental problems. For

example, to help forest management in the Amazon, cultural perspectives of the

Guarayo indigenous people in Bolivia showed that ecological perspectives of sparse

and plentiful species were similar to scientific perspectives of intrinsic growth rate (k or

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r-related species), determined what game species was considered valuable to the

Guarayo community and that subsistence hunting and fishing would continue to be

important. Since the landscape is changing, understanding the indigenous people’s

cultural perspectives is important and fundamental to wildlife management and

sustainability of the Bolivian Amazon natural resources (Van Holt et al. 2010). A study in

Hawai‘i that used CCT revealed a high-yield of similar cultural perspectives among

hand-line fishers and fishery scientists regarding yellowfin tuna stock structure, fish

movements, resource abundance, stock conditions, and fishery interaction (Miller et al.

2004). This information was valuable to the management of Hawai‘i’s yellowfin tuna

fishery because it provided data about the contemporary state of the fishery and

revealed uniform cultural perspectives between hand-line fishers and fishery scientists

(Miller et al. 2004). In North Carolina, cultural perspectives about coastal resource

problems associated with fishing among recreational fishers, commercial fishers, and

coastal resource managers were analyzed with CCT (Johnson and Griffith 2010). Unlike

the yellowfin tuna study in Hawai‘i, this study revealed striking differences in cultural

perspectives between the groups but recreational and commercial fishers did agree on

several underline fishing issues, which may reflect shared values toward the philosophy

and a willingness to support future fishery resource management actions in North

Carolina. This chapter contributes to the building knowledge of jellyfish research by

using CCT to determine cultural perspectives of jellyfish across four groups of people.

By utilizing perspectives of jellyfish-human interactions, these data provides quantitative

social science data that will benefit coastal communities prone to jellyfish occurrences.

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METHODS

Cultural group classification and determination

The cultural groups were classified by the paradigm that cultures form around

specific recreation and leisure activities (McDonough 2013). Under this definition, the

“social world” of each group would have its own unique culture with behavioral norms,

expectations, roles, language, and items such as clothing and gear (Ditton et al. 1992).

The survey’s participants self-identified themselves as fishers (people whose main

career was commercial fishing or hobby was recreational fishing) or coastal

recreationists (people whose main career or hobby entails long periods of time in the

water, such as swimming, surfing, etc.). The coastal and jellyfish researchers did not

self-identify themselves; instead, I identified them by their research interests and

publications via scholarly searches. Although research is usually a career and not

necessarily associated with recreation/leisure, coastal and jellyfish researchers were

expected to have different social worlds and thus were classified as two distinct cultural

groups just as fishers and recreationists.

Study design and sampling framework

I created the jellyfish survey under the guidelines presented by Weller (2007).

The survey contained 64 statements; 32 statements on jellyfish ecology and 32

statements on how jellyfish influence society. The jellyfish survey (Appendix 1) was

distributed in July 2012. All participants were adults (18+ years of age) and U.S.

citizens. For the fishers and coastal recreationists, only NC residents were solicited with

individual face-to-face interviews (N = 75, successful interview = 30 per group) were

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conducted at coastal areas in eastern North Carolina that are in proximity to areas

known to experience jellyfish occurrences (Figure 2). All face-to-face interviews were

conducted within two week period and the fishers and recreationists were found

arbitrarily throughout the 25 areas (Figure 2). Mailed surveys (N = 82 total, returned =

38) were sent to coastal researchers in North Carolina, which consisted of researchers

from academic institutions, federal and state agencies, and non-profit organizations.

Mailed surveys (n= 47, returned = 20) were also sent to jellyfish researchers residing in

the United States. The IRB for this study was obtained from the University & Medical

Center Review Board at East Carolina University, number UMCIRB 12-000609.

Three jellyfish species that are frequently sighted in eastern North Carolina are

the comb jellyfish Mnemiopsis leidyi [A. Agassiz, 1865] estuarine sea nettle Chrysaora

quinquecirrha [Desor, 1848] (Williams and Deubler 1968) and the oceanic cannonball

jellyfish Stomolophus meleagris [L. Agassiz, 1860] (Calder 1982). The sea nettle C.

quinquecirrha is a jellyfish that is known to sting people (Schultz and Cargo 1969) but

the encounters with the comb jellyfish M. leidyi and the cannonball jellyfish S. meleagris

do not typically harm people. Since the severity of jellyfish-human interactions is

dependent on if the jellyfish species’ stings hurt people, I surveyed fishers and

recreationists at estuarine and oceanic areas where potentially harmful and harmless

species are found (Figure 2).

Statistical Analyses

The root of CCT is based the idea that culture is a set of learned and shared

beliefs and behaviors (Weller 2007). Cultural beliefs are affected by the social norms

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(normative beliefs) associated with a group (Heywood 1996; Vaske and Whittaker 2004)

and group culture is the most frequently held items of knowledge and belief (D'Andrade

1987). CCT generates 1) a culturally correct answer key derived from the average of

participants’ responses and 2) cultural competency scores or cultural competencies.

Cultural competency, or cultural expertise of an individual, is calculated by comparing

agreement between the study’s participants and in general, higher values equate to

greater cultural competencies (Weller 2007). To analyze cultural competencies within

and across groups, a factor analysis of an informant by informant agreement matrix was

performed (D'Andrade 1987; Vaske and Whittaker 2004) on all participant’s responses

and the participant responses separated by their group. The participants are considered

to have a consensus about the domain analyzed if the 1st and 2nd factor loading (ratio) is

greater than 3, there are no negative competency values, and there is a high amount of

agreement in responses among participants (Romney et al. 1986; Weller 2007). Cultural

competency of each participant was calculated by comparing agreement between all

pairs of the survey’s participants (Weller 2007) and represents the first factor loading of

the factor analysis (Romney et al. 1986). The statistical program UCINET (version

6.322) from Analytic Technologies was used to perform all factor analyses and generate

cultural competencies. Mean cultural competencies of all participants separated by their

group were compared with a one-way ANOVA. Variance of the ANOVA was further

tested for homogeneity or heterogeneity with a Levene test of variance. To determine

how mean cultural competency differed between groups, a Tukey post-hoc test was

performed. Comparison of the jellyfish survey’s culturally correct answers for each

statement derived from the each groups’ consensus analysis were classified according

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to accordance and discordance, where “accordance” refers to the unanimous

agreement of culturally correct answers among the four groups and “discordance” refers

to at least one of the four groups disagreed. To visualize participant agreement within

and across groups, UCINET was used to create metric multidimensional scaling

diagrams based on the aggregate proximity matrix generated from the factor analysis.

RESULTS

Consensus analyses and cultural competencies of jellyfish perception

The consensus analyses performed on all of the statements within the jellyfish

survey for all of the participants (N = 118) and each group had a first to second

eigenvalue ratio greater than 3 and no negative values. Of the groups, fishers (ratio =

3.39, SD = 0.07) and recreationists (ratio = 4.55, SD = 0.06) showed more intragroup

variation in jellyfish perception than the coastal researchers (ratio = 5.58, SD = 0.04)

and jellyfish researchers (ratio = 5.63, SD = 0.06) (Table 1). The ANOVA revealed that

mean cultural competencies across all groups were significantly different (F-value =

7.97, p-value = 7.21e-05) and the Levene test of variance confirmed that variance was

homogeneous among the competency values (Test statistic = 1.43, p-value = 0.24).

Therefore, mean cultural competencies were similar between fishers (0.23) and

recreationists (0.24) (Tukey p-value = 1.00), fishers (0.23) and coastal researchers

(0.27) (Tukey p-value = 0.05), recreationists (0.24) and coastal researchers (0.27)

(Tukey p-value = 0.09), and coastal researchers (0.27) and jellyfish researchers (0.30)

(Tukey p-value = 0.15). Significant differences in mean cultural competency were noted

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between the fishers (0.23) and jellyfish researchers (0.30) (Tukey p-value = 0.00) and

recreationists (0.24) and jellyfish researchers (0.30) (Tukey p-value = 0.00) (Figure 3).

Accordance of jellyfish survey ecological statements

The culturally correct answers to the ecological statements of each group were

often in accordance. The groups agreed that statements such as jellyfish are animals

(#1), live longer than a year (#3), jellyfish eat fish and shrimp (#14, #15), jellyfish are

able to eat more than it’s body weight (#17) and all groups agreed that turtles and fish

eat jellyfish (#20, #22). In addition, false statements, myths, and misunderstandings

about jellyfish ecology were accurately identified by answer accordance regarding

jellyfish ecology and stings. These statements included jellyfish use fins to swim (#7),

jellyfish can swim against moving water (#9), jellyfish need to surface to breathe and eat

(#12, #13), jellyfish do not need to eat to survive (#18), jellyfish are top trophic level

organisms in food webs like sharks (#23, Figure 4A), jellyfish are low trophic level

organisms in food webs like plants (#26, Figure 4D), and jellyfish do not sting to

reproduce (#29), communicate (#30), and can sting more than once (#31) (Table 2).

Accordance of jellyfish survey societal statements

The culturally correct answers to the societal statements of each group were also

in frequent accordance. Regarding jellyfish stings, answers from all groups accurately

identified that jellyfish stings are felt on a person’s body (#35) and ‘vinegar is the best

remedy for a jellyfish sting’ (#38) was mutually agreed upon (Table 3). Several answers

to the myths of jellyfish stings were also in accordance; for example, red-color jellyfish

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stings hurt more than other-colored jellyfish stings (#33), jellyfish cannot sting the palm

of a person’s hand (#34), and jellyfish do not need to touch a person to sting them

(#37). Accordance of answers to the perception of jellyfish and society showed that all

groups agreed that people would not benefit if there were no jellyfish (#47) and jellyfish

have economic value (#48). However, all groups also agreed that the presence of

jellyfish will deter people from water activities (#53). When asked ‘how many jellyfish

seen in water will make a person stop their water activities,’ there was accordance that

water activities will continue if there are fewer than five jellyfish seen in water (#41).

Hearing that someone else was stung by jellyfish will not deter people from doing water

activities (#57) but if people are stung by a jellyfish they will stop their water activities

(#58). The presence of jellyfish affects vacationing (#52) but there was accordance that

people will take a vacation to areas where jellyfish are sighted regularly (#49), and

people will book a return vacation to a destination even if they are stung by jellyfish

(#50). For recreation, group accordance revealed that if jellyfish are always present in

an area, people will find another place for recreation (#56). When asked if jellyfish

appear without warning and if people know where jellyfish come from, all groups agreed

that people think jellyfish appear without warning (#59) and that people do not know

where jellyfish come from (#60). Incidentally, all groups disagreed with the statement

that jellyfish are taking over aquatic ecosystems worldwide (#62) (Table 3).

Discordance of jellyfish survey ecological statements

Answers to 14 ecological statements showed discordance among group

responses, possibly reflecting expertise regarding jellyfish biology and ecology (Table

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4). When specific questions were asked about jellyfish, the jellyfish researchers agreed

with the statements jellyfish have eyes (#4), no brain (#5), no heart (#6), and do not use

tentacles to swim (#8), which are also biologically correct. The fishers, recreationists,

and coastal researchers varied in their answers to these statements but when

compared to the jellyfish researchers, coastal researchers agreed that jellyfish do not

have a brain (#5) and a heart (#6) and these answers are biologically correct. Jellyfish

thrive in murky water (#10) and when asked, jellyfish researchers agreed and the other

groups disagreed. Jellyfish also do not need a lot of air-in-water to survive (#11) and all

groups except the coastal researchers agreed. Some of the false statements that

inquired about what jellyfish consume and what eats jellyfish (#16, #19 and #21) varied

in group response, specifically fishers agreed that jellyfish eat plants (#16) and dolphins

eat jellyfish (#21), which is biologically incorrect. Jellyfish researchers answered that

jellyfish are mid-trophic level organisms like fish (#24, Figure 4B) and the current

research regarding jellyfish in food webs indicates that this is the correct answer

(Brodeur et al. 2011; Purcell and Decker 2005; Suchman et al. 2008), whereas fishers

and recreationist responses suggested that jellyfish are lower trophic level organisms

like shrimp (#25, Figure 4C). As a group, the coastal researchers did not select any of

the food web pictures but when individual participant answers were analyzed the

response from coastal researchers varied. Forty-percent of coastal researchers

selected the same food web picture as fishers and recreationists (#25, Figure 4C) and

29% of coastal researchers selected the same food web picture as the jellyfish

researchers (#24, Figure 4B). The remaining 31% coastal researchers did not select

any of the food webs pictures. For the sting questions, only jellyfish researchers agreed

129

that all jellyfish sting (#27). Moreover, statements that reported myths about jellyfish

stings were also highly variable in group response, where fishers and recreationists

agreed that jellyfish sting to protect themselves (#28) but coastal and jellyfish

researchers disagreed. Lastly, fishers were the only group that agreed to the statement

“jellyfish cannot control their sting” (#32) (Table 4).

Discordance of jellyfish survey societal statements

There were nine societal statements that were in discordance across all groups.

For example, the jellyfish researchers disagreed with the statement ‘jellyfish cannot

sting a person through their clothes’ (#36) and fishers were the only group that agreed

to the statement that urine as the best remedy for a jellyfish sting (#39). When asked

‘how many jellyfish seen in water will make a person stop their water activities,’ there

were different answers among the groups. Coastal researchers and jellyfish researchers

agreed that seeing five or more jellyfish in water will make people stop water activities

(#44) but fishers and recreationists disagreed. Moreover, fishers were the only group

that agreed with the statement ‘people will continue their regular water activities

(hobbies) if jellyfish are in the water’ (#54). When asked if people would book a return

vacation to an area if they were stung by jellyfish (#51), recreationists and jellyfish

researchers agreed but fishers and coastal researchers disagreed. Only coastal

researchers did not select the statement that people will leave a recreation area if

jellyfish are present (#55). When asked if there is a way to predict when large numbers

of jellyfish will appear (#61), fishers and coastal researchers disagreed whereas

recreationists and jellyfish researchers agreed. Lastly, when asked if jellyfish are

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frequently reported in the media (newspapers, websites, etc.) (#63, #64), only jellyfish

researchers agreed (Table 5).

MDS of ecological statements

For the ecological statements, the metric MDS showed higher agreement among

the jellyfish researchers than other groups. The jellyfish researchers are close to the

coastal researchers but the recreationists and fishers are quite dispersed from both

coastal and jellyfish researchers. The most dispersed agreement was noted in the

fishers. However, it should be noted that agreement among the fishers and

recreationists is equally distant from the coastal and jellyfish researchers (Figure 4).

MDS of societal statements

For the societal statements, there was varying amounts of agreement across all

groups with no cluster as in the ecological statements (Figure 4). This suggests that all

participants of the jellyfish survey did not share the same agreement toward the cultural

statements concerning jellyfish and society (Figure 5).

DISCUSSION

I found that perspectives of jellyfish ecology varied among the groups but societal

perspectives of jellyfish were similar. The cultural perspectives of jellyfish researchers

were closely aligned with what is biologically known about jellyfish, but perspectives on

jellyfish ecology among fishers, recreationists and coastal researchers were less clear.

All group perspectives on the influence of jellyfish on water activities showed that

131

although jellyfish may be problematic to coastal communities, water activities and

associated revenue may not be affected due to varying amounts of tolerance toward

seeing jellyfish in water and stings.

Cultural consensus on jellyfish ecology

The results of the jellyfish survey’s ecological statements support my first

hypothesis that the cultural groups would share similar cultural perspectives of jellyfish

ecology, with the obvious exception of jellyfish researchers. Mean cultural competencies

of coastal and jellyfish researchers were similar but the range of cultural competency

among the coastal researchers overlapped with fishers and recreationists more than

jellyfish researchers (Figure 3). Most of the culturally correct answer keys for the

ecological statements regarding jellyfish in food webs were in accordance across all

groups (Table 2). Furthermore, all groups accurately selected what is scientifically

known about jellyfish as predators and prey. However, the overlap in cultural

competency and similar mean cultural competencies between fishers, recreationists,

and coastal researchers (Figure 3) suggests that knowledge of jellyfish in ecosystems is

less clear at both the public and research interfaces.

The statements regarding the feeding ecology of jellyfish had culturally correct

answer keys in discordance (Table 4). Specifically, there seems to be confusion on the

role jellyfish play in food webs (#23-26, Table 2 & 4). Fishers, recreationists, and most

of the coastal researchers (41%) believed that jellyfish in nature are best described as

an organism that shares a similar trophic level as shrimp (Figure 4C). Since the current

data on jellyfish in food webs has shown that jellyfish are intermediate to top level

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trophic level predators (Figure 4B) (Brodeur et al. 2011; Mills 1995; Purcell et al. 2007)

and jellyfish researchers selected this food web picture as the culturally correct answer,

I can infer that fishers, recreationists, and 39% of coastal researchers all selected the

incorrect food web picture that best describes jellyfish in food webs. It should be noted

that 31% of coastal researchers did not select a food web picture and of the 31% only

1% wrote comments about the food web pictures being inaccurate due to the energy

flow or arrow direction pointing downwards instead of upwards and the food web

diagrams were too simplistic for them to make a choice. Although the simple food web

pictures used in the jellyfish survey were created for the purposes of easy interpretation

across all participation groups, I felt that the pictures were adequate enough to visualize

where jellyfish belong in food webs. Moreover, 29% of coastal researchers did select

the culturally and scientifically correct food web picture, which demonstrates that some

of the coastal researchers surveyed were familiar with the role jellyfish play in foods

webs.

It is plausible that selecting food web concepts by reading statements versus

selecting a food web picture may have been easier for the survey participants to

comprehend. This ideology may reflect what scientific educators refer to as an

“ecological misconception” (Cherrett 1989). Research has shown that students (4th

grade to college level juniors and seniors) comprehend and internalize food chain

concepts (what eats what), but have difficulties when food chain principles are applied

to the complexities of food web dynamics (Adeniyi 1985; Griffiths and Grant 1985;

Munson 1994). Ecological misconceptions are typically formed by students who utilize

their prior knowledge and experience when asked about scientific phenomenon

133

(Hewson and Hewson 1988; Posner et al. 1982). Moreover, ecological misconceptions

are created when students have an incorrect interpretation or hold an alternative

understanding of the subject matter (Munson 1994). Since the ecological statements

read in a systematic format (Table 2, #14, #15, #20 and #22), the information processed

by the participant could have been ‘chain-like’ or ordinal, where the participant could

internalize the information easier than applying this ecological knowledge to the food

web pictures. Also, if the incorrect food web choice was indeed an ecological

misconception, which stems from the participant’s prior knowledge and experience with

jellyfish, it is conceivable that the lack of education and data about jellyfish in

ecosystems may explain why fishers, recreationists, and coastal researchers placed

jellyfish improperly in food webs.

Cultural consensus on the societal role of jellyfish

The results of the jellyfish survey’s societal statements supported my 2nd

hypothesis that all groups would have similar cultural perspectives of how jellyfish

influence society (Table 3). The MDS of the jellyfish survey’s societal statements did not

show a clear distinction or cluster of a group’s agreement (Figure 5). Instead

participants from all groups were randomly distributed throughout the matrix and varied

spatially from each other. This showed that, unlike the jellyfish survey’s ecological

statements, there was less agreement within each group in the statements regarding

how jellyfish influence society.

I found that the cultural perspectives of how jellyfish influence water activities,

recreation, and vacationing differed among all groups. Although all groups agreed that

134

seeing fewer than five jellyfish would not stop people from engaging in water activities

(Table 3, #41), there were different perspectives in the termination of water activities of

seeing more than five jellyfish in water (Table 5, #44). Specifically, fishers and

recreationists felt that seeing more than five jellyfish in the water would not stop water

activities and coastal and jellyfish researchers thought water activities would cease.

Since the survey was distributed in different coastal recreation areas in eastern North

Carolina (Figure 1), it is possible that the jellyfish species that the fishers or

recreationists were thinking about while taking the survey, have frequently observed,

and/or had personal experiences with are not species that is harmful to humans. Of the

three jellyfish species that are sighted frequently the comb jellyfish Mnemiopsis leidyi [A.

Agassiz, 1865] and the oceanic cannonball jellyfish Stomolophus meleagris [L. Agassiz,

1860] are usually not harmful to humans but the sea nettle C. quinquecirrha is a jellyfish

that is known to cause painful stings (Schultz and Cargo 1969). Twenty-three percent of

fishers and 37% of recreationists felt that water activities would continue with more than

five jellyfish conduct their water activities in oceanic areas where the relatively harmless

cannonball jellyfish Stomolophus meleagris are observed. Continuing water activities

with more than five jellyfish may also coincide with the type of protective equipment

fishers and recreationists used when engaging in their water activities. Fishers in North

Carolina are known to use waders and some of the recreationists wear wetsuits, which

prevent contact with jellyfish. In fact, among the recreationists that enjoy water activities

in estuarine areas where stinging sea nettles C. quinquecirrha are abundant were

analyzed, only 2% felt that seeing more than five jellyfish would stop water activities. If

the sea nettle C. quinquecirrha was the jellyfish that were frequently in contact with

135

these recreationists, it could be that these individuals are more tolerant of sea nettle C.

quinquecirrha stings. Conversely, although the consensus analysis showed that the

fishers agreed with the recreationists, 46% of fishers who frequently fished in areas

where sea nettles C. quinquecirrha are abundant felt that seeing more than five jellyfish

would stop water activities. However, despite the negative repercussions of jellyfish

stings, all groups agreed that people would take a vacation to areas where jellyfish are

sighted regularly even if they are stung by jellyfish.

All groups felt that jellyfish have economic value. From a wildlife management

standpoint, wildlife has either a positive or negative social value dependent on human

perspective (Brown and Manfredo 1987). With this in mind, coastal communities may

revisit their perspectives of jellyfish depending on changing environmental-social

pressures. For example, the cannonball jellyfish S. meleagris is an abundant jellyfish

species along the southeastern and Gulf coasts of the U.S. and large populations are

observed off the Georgia coast from March to October (Calder 1982). According to a

local newspaper, in 1998 the harvests of the cannonball jellyfish S. meleagris turned

Georgia’s shrimping industry into the third-largest commercial fishery by jellyfish weight

alone. Shrimpers can earn $0.06 per pound of jellyfish and the processing plant can

process 60,000 pounds at a time. Due to high fuel costs, some shrimpers have

completely switched from harvesting shrimp to jellyfish and all jellyfish harvests are

exported directly to Asia (Landers 2011). It is unknown how this jellyfish fishery affects

natural populations of jellyfish, but researchers have reported problems with the

cannonball jellyfish S. meleagris preventing shrimping activities in Georgia and South

Carolina for decades (Jenkins 2012; Kraeuter and Setzler 1975). Georgia’s cannonball

136

fishery could be an example of how social value of jellyfish has changed overtime from

negative to positive social value thereby altering public perception of jellyfish in the

United States.

Implications for managing jellyfish-human interactions

These data, along with other research on perspectives of the jellyfish in society,

will be useful for economic studies of tourism areas that are known to experience

jellyfish occurrences and coastal managers to help mitigate deleterious jellyfish-human

interactions. For example, the use of barrier nets to designate swimming areas in

coastal areas prone to jellyfish occurrences may reduce the number of jellyfish to the

“fewer than five” allowance of jellyfish sighted. Since seeing fewer than five jellyfish will

not stop water activities, negative effects on coastal recreation could be minimized. Low

jellyfish numbers in swimming areas will also reduce the potential of jellyfish stings.

Thus understanding the cultural perspective of “fewer than five” could help with coastal

recreation and subsequently tourism in North Carolina, and if the cultural perspectives

are similar, other U.S. coastal states where jellyfish may be problematic. On the other

hand, if cultural perspectives are different, there may be instances where more jellyfish

sighted in recreation areas may not affect water activities. For example, in a recent

study of jellyfish public perception on German beaches, beach users were reported to

tolerate periodic and moderately increasing jellyfish numbers in the water (Baumann

and Schernewski 2012). Additionally, Baumann and Schernewski’s (2012) research

investigated “willingness to pay” for a swimming area that is free from jellyfish and found

that most beach users would not pay more for a netted swimming area devoid of

137

jellyfish. However, the author’s noted that the public is well-informed that there are no

“life-threatening jellyfish species” in this area (Baumann and Schernewski 2012).

Moreover, information (warning signs, information panels and pamphlets) on jellyfish

that is readily available on-site proved to help beach users accept and feel better about

jellyfish being on German beaches (Baumann and Schernewski 2012). Although our

methodologies were different, this study and my cultural consensus analysis show how

the utility of jellyfish public perspective can be used to evaluate the extent of jellyfish-

human interactions.

CONCLUSIONS

Public perspective of marine predators, like jellyfish, has substantial effects on

scientific explorations as well as the utilization of coastal recreation areas, and

subsequently, ecosystem services and economics. For example, like jellyfish, the

presence of sharks has also influenced coastal recreation because humans have feared

sharks for centuries and shark attacks have been detrimental to tourism (Cliff 1991).

Early shark research in the 1960s and 1970s focused on shark physiology and

behavior, including attack behavior, with the goal of human protection from sharks.

Overtime, shark research changed focus when shark-control programs proved to be

more effective in mitigating shark attacks, but during this investigation process a

plethora of life history research on sharks was obtained (Simpfendorfer et al. 2011).

More shark research equated to a further understanding of the role of sharks in

ecosystems and over time public perception has shifted from “adventure-seeking

hunters” to “nature-seeking observers” of sharks (Whatmough et al. 2011). Today,

138

instead of exploring ways to protect humans from sharks, shark conservation and

management are at the forefront of research endeavors (Dulvy et al. 2008;

Simpfendorfer et al. 2011). Although conserving jellyfish is an unprecedented scientific

proposition, it is possible that jellyfish research may follow similar scientific exploration

paths as shark research, especially since jellyfish have significant effects on water

activities, recreation and vacationing. Jellyfish researchers are hopeful that jellyfish

research continues to gain more attention as the scientific community accepts

ecosystem-based fishery management and trans-disciplinary science, which includes

both ecological and societal research questions and approaches to promote

environmental sustainability (Condon et al. 2012; Lang et al. 2012; Pikitch et al. 2004;

Purcell 2012). In time, as more research is conducted on jellyfish and knowledge is

iterated to the public through different media outlets, education, and accurate scientific

reporting, public perspective of jellyfish will change and the cultural knowledge of

jellyfish in ecosystems are likely to improve.

139

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Table 1. Results of the consensus analyses showing mean, standard deviation, and

minimum and maximum cultural competency values (first factor scores) for each run of

the analysis, with the ratio of the first to second eigenvalues.

Table 2. Ecological statements that all groups were in accordance, with culturally

correct answers shown (1 = agree, 0 = disagree; F = fishers, R = recreationists, CR =

coastal researchers, JR = jellyfish researchers).

Table 3. Societal statements that all groups were in accordance, with culturally correct

answers shown (1 = agree, 0 = disagree; F = fishers, R = recreationists, CR = coastal

researchers, JR = jellyfish researchers).

Table 4. Ecological statements that all groups were in discordance, with culturally

correct answers shown (1 = agree, 0 = disagree; F = fishers, R = recreationists, CR =

coastal researchers, JR = jellyfish researchers).

Table 5. Societal statements that all groups were in discordance, with culturally correct

answers shown (1 = agree, 0 = disagree; F = fishers, R = recreationists, CR = coastal

researchers, JR = jellyfish researchers).

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Figure 1. Results from an ISI Web of Science search for the terms estuary +

phytoplankton + zooplankton + fish and + jellyfish. The term ‘jellyfish’ included jellyfish,

gelatinous, ctenophore and sea nettle.

Figure 2. Face to face interviews were conducted at coastal areas known to experience

jellyfish occurrences in eastern North Carolina (circles).

Figure 3. Boxplots of cultural competency derived from the consensus analysis of

jellyfish survey. The same letter indicates similar mean cultural competencies across

groups. Circles indicate outliers > 1.5 times of lower quartile and horizontal bars within

the box plots indicate the cultural competency median. The upper box above the

median extends to the third quartile and the lower box extends to the first quartile. The

bars extend to the largest and smallest values, respectively.

Figure 4. All survey participants were asked to circle the food web picture(s) that best

described jellyfish in food webs (survey questions #23 and #26, Table 2; #24 and #25,

Table 4).

Figure 5. Metric multidimensional scaling of the ecological statements agreement

derived from the aggregate proximity matrix of the consensus analysis (stress = 0.329),

where circles = fishers (N = 30), triangles = recreationists (N = 30), squares = coastal

researchers (N = 38) and diamonds = jellyfish researchers (N = 20).

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Figure 6. Metric multidimensional scaling of the societal statements agreement derived

from the aggregate proximity matrix of the consensus analysis (stress = 0.314), where

circles = fishers (N = 30), triangles = recreationists (N = 30), squares = coastal

researchers (N = 38) and diamonds = jellyfish researchers (N = 20).

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Table 1.

All Participants (N = 118)

Fishers (N = 30)

Recreationists (N = 30)

Coastal

Researchers (N = 38)

Jellyfish

Researchers (N = 20)

Mean 0.25 0.23 0.07 0.02 0.37 3.39

0.24 0.06 0.01 0.34 4.55

0.27 0.04 0.18 0.35 5.58

0.30 0.06 0.18 0.40 5.63

SD 0.06 Min 0.03 Max 0.35 Ratio 4.39

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Table 2. # Ecological Statements F R CR JR 1 Jellyfish are animals 1 1 1 1 2 Jellyfish are plants 0 0 0 0 3 Jellyfish live longer than a year 1 1 1 1 7 Jellyfish use fins to swim 0 0 0 0 9 Jellyfish can swim against moving water 0 0 0 0 12 Jellyfish need to come up to the surface to breathe 0 0 0 0 13 Jellyfish need to come up to the surface to eat 0 0 0 0 14 Jellyfish eat fish 1 1 1 1 15 Jellyfish eat shrimp 1 1 1 1 17 Jellyfish can eat more than their body weight 1 1 1 1 18 Jellyfish do not need to eat to survive 0 0 0 0 20 Turtles eat jellyfish 1 1 1 1 22 Fish eat jellyfish 1 1 1 1 23 The food web picture that illustrates that jellyfish are at

the same trophic level as sharks best describes jellyfish in nature

0 0 0 0

26 The food web picture that illustrates that jellyfish are at the same trophic level as plants best describes jellyfish in nature

0 0 0 0

29 Jellyfish sting to reproduce 0 0 0 0 30 Jellyfish sting to communicate 0 0 0 0 31 A jellyfish can sting only once 0 0 0 0

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Table 3. # Societal Statements F R CR JR 33 Stings from red-colored jellyfish hurt more than other-

colored jellyfish 0 0 0 0

34 Jellyfish cannot sting the palm of a person’s hand 0 0 0 0 35 Jellyfish stings are felt on a person’s body 1 1 1 1 37 Jellyfish do not need to touch a person to sting them 0 0 0 0 38 Vinegar is the best remedy for a jellyfish sting 1 1 1 1 40 There is no remedy for a jellyfish sting 0 0 0 0 41 Seeing fewer than five jellyfish in water will make people

stop their water activities. 0 0 0 0

47 People would benefit if there were no jellyfish 0 0 0 0 48 Jellyfish have economic value 1 1 1 1 49 People will not take a vacation to areas where jellyfish

are sighted regularly 0 0 0 0

50 People will not book a return vacation to an area if they were stung by jellyfish

0 0 0 0

52 The presence of jellyfish does not affect vacationing 0 0 0 0 53 The presence of jellyfish will deter people from doing

water activities 1 1 1 1

56 If jellyfish are always present in an area, people will find another place for recreation

1 1 1 1

57 If people hear that someone was stung by jellyfish they will stop their water activities

0 0 0 0

58 If people are stung by jellyfish, they will stop their water activities

1 1 1 1

59 People think jellyfish will appear without warning 1 1 1 1 60 People know where jellyfish come from 0 0 0 0 62 Jellyfish are taking over aquatic ecosystems worldwide 0 0 0 0

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Table 4. # Ecological Statements F R CR JR 4 Jellyfish have eyes 0 0 0 1 5 Jellyfish have a brain 1 1 0 0 6 Jellyfish have a heart 1 1 0 0 8 Jellyfish use tentacles to swim 1 1 0 0 10 Jellyfish thrive in murky water 0 0 0 1 11 Jellyfish do not need a lot of air-in-water to survive 1 1 0 1 16 Jellyfish eat plants 1 0 0 0 19 People eat jellyfish 0 0 1 1 21 Dolphins eat jellyfish 1 0 0 0 24 The food web picture that illustrates that jellyfish are at

the same trophic level as fish best describes jellyfish in nature

0 0 0 1

25 The food web picture that illustrates that jellyfish are at the same trophic level as shrimp best describes jellyfish in nature

1 1 0 0

27 All jellyfish sting 0 0 0 1 28 Jellyfish sting to protect themselves 1 1 0 0 32 Jellyfish cannot control their sting 0 1 1 1

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Table 5.

# Societal Statements F R CR JR 36 Jellyfish cannot sting a person through their clothes 0 0 0 1 39 Urine is the best remedy for a jellyfish sting 1 0 0 0

44 Seeing more than five jellyfish in water will make people stop their water activities

0 0 1 1

51 People will not book a return vacation to an area if they were stung by jellyfish

0 1 0 1

54 People will continue their regular water activities (hobbies) if jellyfish are in the water

1 0 0 0

55 People will leave a recreation area if jellyfish are present 1 1 0 1 61 There is no way to predict when large numbers of

jellyfish will appear 1 0 1 0

63 Jellyfish are frequently reported in the media (newspapers, websites, etc.)

0 0 0 1

64 Jellyfish are rarely reported in the media (newspapers, websites, etc.)

1 1 1 0

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Figure 1.

161

Figure 2.

162

Figure 3.

163

Figure 4.

164

Figure 5.

165

Figure 6.

166

Appendix 1.

For these questions, please tell us if people agree or disagree with the following statements about jellyfish.

01. Jellyfish are animals Agree Disagree

02. Jellyfish are plants Agree Disagree

03. Jellyfish live longer than a year Agree Disagree

04. Jellyfish have eyes Agree Disagree

05. Jellyfish have a brain Agree Disagree

06. Jellyfish have a heart Agree Disagree

07. Jellyfish use fins to swim Agree Disagree

08. Jellyfish use tentacles to swim Agree Disagree

09. Jellyfish can swim against moving water Agree Disagree

10. Jellyfish thrive in murky water Agree Disagree

11. Jellyfish do not need a lot of air-in-water to survive Agree Disagree

12. Jellyfish need to come up to the surface to breathe Agree Disagree

13. Jellyfish come up to the surface to eat Agree Disagree

14. Jellyfish eat fish Agree Disagree

15. Jellyfish eat shrimp Agree Disagree

16. Jellyfish eat plants Agree Disagree

17. Jellyfish can eat more than their body weight Agree Disagree

18. Jellyfish do not need to eat to survive Agree Disagree

19. People eat jellyfish Agree Disagree

20. Turtles eat jellyfish Agree Disagree

21. Dolphins eat jellyfish Agree Disagree

22. Fish eat jellyfish Agree Disagree

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Which food web picture or pictures best describes jellyfish in nature? Please circle your answer(s).

For these questions, please tell us if people agree or disagree with the following statements about jellyfish stings.

01. All jellyfish sting Agree Disagree

02. Jellyfish sting to protect themselves Agree Disagree

03. Jellyfish sting to reproduce Agree Disagree

04. Jellyfish sting to communicate Agree Disagree

05. A jellyfish can sting only once Agree Disagree

06. Jellyfish cannot control their sting Agree Disagree

07. Stings from red-colored jellyfish hurt more than other-colored jellyfish Agree Disagree

08. Jellyfish cannot sting the palm of a person’s hand Agree Disagree

09. Jellyfish stings are felt on a person’s body Agree Disagree

10. Jellyfish cannot sting a person through their clothes Agree Disagree

11. Jellyfish do not need to touch a person to sting them Agree Disagree

12. Vinegar is the best remedy for a jellyfish sting Agree Disagree

13. Urine is the best remedy for a jellyfish sting Agree Disagree

14. There is no remedy for a jellyfish sting Agree Disagree

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How many jellyfish seen in the water will make people stop their water activities? Please circle your answer.

0 Jellyfish 1 Jellyfish



169

For these questions, think about jellyfish and society. Please tell us if people agree or disagree with the following statements.

01. People would benefit if there were no jellyfish Agree Disagree

02. Jellyfish have economic value Agree Disagree

03. People will not take a vacation to areas were jellyfish are sighted regularly Agree Disagree

04. People will not book a return vacation to an area if they saw jellyfish in the water Agree Disagree

05. People will not book a return vacation to an area if they were stung by jellyfish Agree Disagree

06. The presence of jellyfish does not affect vacationing Agree Disagree

07. The presence of jellyfish will deter people from doing water activities Agree Disagree

08. People will continue their regular water activities (hobbies) if jellyfish are in the water Agree Disagree

09. People will leave a recreation area if jellyfish are present Agree Disagree

10. If jellyfish are always present at an area, people will find another place for recreation Agree Disagree

11. If people hear that someone was stung by jellyfish they will stop water activities Agree Disagree

12. If people are stung by jellyfish, they will stop water activities Agree Disagree

13. Jellyfish will appear in large numbers without warning Agree Disagree

14. People know where jellyfish come from Agree Disagree

15. There is no way to predict when large numbers of jellyfish will appear Agree Disagree

16. Jellyfish are taking over aquatic ecosystems worldwide Agree Disagree

17. Jellyfish are frequently reported in the media (newspapers, websites, etc.) Agree Disagree

18. Jellyfish are rarely reported in the media (newspapers, websites, etc.) Agree Disagree

CHAPTER 5

Dissertation conclusions and management implications

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Jellyfish-human interactions in North Carolina

The overall objective of my dissertation was to identify physical, ecological and

social drivers of jellyfish-human interactions in North Carolina. Since large numbers of

jellyfish have created many ecological and societal effects on coastal communities, I

chose a multifaceted research approach to gather physical and ecological data on what

influences a harmful jellyfish species in a highly-used estuarine recreation area and to

evaluate this interaction as well as other jellyfish-human interactions in North Carolina.

By studying jellyfish-human interactions, my dissertation has provided data on jellyfish

in North Carolina and this knowledge will help coastal management with future

ecological and socio-economic issues caused by jellyfish.

I found that the distribution and abundance of C. quinquecirrha at six heavily

used recreation sites was related to wind events (3-8 m s-1 at Cherry Point Marine

Weather Station and 2-7 m s-1 at Cape Hatteras Weather Station). The dominant wind

directions during the spring and summer in the months associated with C. quinquecirrha

occurrences in 2011 and 2012 were SSE-SSW and these wind directions generate ~10

cm s-1 water currents that were likely to move C. quinquecirrha along and across the

NRE estuarine shorelines. The highest abundance (m2) was observed on the South

shoreline, specifically at PC and SF, which are coastline sites located at NRE bend. It is

possible that these sites experienced the highest C. quinquecirrha because of the

shallow nature of the area and geomorphology of the NRE thereby making

accumulation of jellyfish more likely than the North shoreline pier sites. When wind

events occurred, I found that SSE to SSW wind directions one day prior to observations

could be correlated with increased C. quinquecirrha abundance along the South

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shoreline and 5 days prior to observations could be correlated with increases on the

North shoreline. Since no other abiotic variables could be correlated with C.

quinquecirrha abundance, my results in chapter 2 indicated that one of the primary

physical drivers of jellyfish-human interactions in the NRE are wind events.

The jellyfish-human interactions of C. quinquecirrha and NRE estuarine

recreation enthusiasts may be manageable with the knowledge that wind speed and

direction is correlated to higher abundances on the North and South shorelines. Future

research should focus on creating a forecast system or adding potential C.

quinquecirrha occurrence to the RENCI ADCIRC storm surge and coastal threats

model. In the interim, readily available wind data via web-based and personal device

social media from services such as the National Weather Service, NOAA, Weather bug

©, Earth Networks, and Wind Alert ©, Weather Flow, Inc. will allow NRE fishers and

recreationists to plan their activities accordingly. This insight may help with mitigating

the potential loss of recreation and tourism revenue as well as provide peace of mind to

people that have trepidation towards jellyfish stings. For example, public

announcements of the likelihood of encountering the stinging box jellyfish A. moseri at

Waikīkī Beach eight to twelve days after a full moon has helped residents and tourists in

Hawai‘i plan their beach-going activities (Thomas et al. 2001). On-site first aid or

outreach education has helped people cope with large amounts of stinging jellyfish in

Australia (Gershwin et al. 2010) and Germany (Baumann and Schernewski 2012).

While conducting field research in the NRE, I noticed that people were very receptive to

learning about jellyfish and because we would have vinegar to treat ourselves if stung

by the sea nettles C. quinquecirrha, we often shared vinegar with people who were also

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stung. After a vinegar treatment, even a most terrified child that experienced a jellyfish

sting did not hesitate to jump back into the water. Since public education and

information about jellyfish has helped mitigate unwanted jellyfish-human interactions, I

recommend the use of signage to help the public know when jellyfish are likely to occur

in large numbers used by the National Park Service within the Neuse River Recreation

Area, Croatan National Forest and private vendors such as the YMCA camps.

Information on jellyfish stings would also benefit the public.

Excluding my dissertation research, there have been three publications that

mention jellyfish in North Carolina (Mallin 1991; Miller 1974; Williams and Deubler 1968)

and of the three, only one investigated jellyfish abundance in the PRE (Miller 1974)

(Figure 1). The use of citizen-science monitoring could determine the abundance and

distribution of jellyfish species, as my visual observations of C. quinquecirrha ~ 2 m2

were similar to Miller’s (1974). Also, traditional net survey techniques (Tucker trawls,

plankton tows, and beach seines) throughout APES could determine spatial and

temporal jellyfish patterns over longer periods of time. C. quinquecirrha predation plays

a critical role in Chesapeake Bay food webs but it is unknown how this estuarine

predator influences the APES ecosystem. Therefore, food web studies in APES should

include C. quinquecirrha and other jellyfish species to assure accurate

conceptualization of trophic pathways.

Although my research in chapter 2 showed no significant correlations with abiotic

variables known to influence C. quinquecirrha; temperature, salinity, and dissolved

oxygen may influence some of the life history stages of the sea nettle, as documented

in Chesapeake Bay (Calder 1974; Condon et al. 2001; Decker et al. 2007; Purcell and

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Decker 2005). For example, it is well known that temperature and salinity affect the

polyp and strobila stages of C. quinquecirrha (Calder 1974; Cargo and Rabenold 1980).

In addition, jellyfish polyps and medusae are more resilient to hypoxia or low dissolved

oxygen than other aquatic species (Condon et al. 2001; Purcell et al. 2001; Thuesen et

al. 2005) and may benefit ecologically by consuming prey in hypoxic areas (Shoji et al.

2010). The tributaries in APES are prone to hypoxia (Paerl et al. 1998; Stanley and

Nixon 1992) and could serve as an important study site for future hypoxia-jellyfish

studies. Additionally, investigations of the abiotic drivers and substrate preferences of

the early life history stages of C. quinquecirrha are essential to determining this species’

vitality in APES and the annual occurrences observed in the NRE and other tributaries.

Although C. quinquecirrha abundances were correlated with wind events and

associated wind-driven circulation, I cannot conclude that the annual occurrence of C.

quinquecirrha is related to sexual reproduction. Instead, my results in chapter 3

indicated that there is potential for sexual reproduction to occur, but there was equal

evidence that sexual reproduction may not be responsible for the annual occurrences.

Moreover, the wind events, discussed in chapter 2, do not appear to aggregate jellyfish

that are sexually reproducing based on my ANOVA analyses of egg diameters. Larger

egg diameters were observed along the South shoreline; however, none of these eggs

were considered mature based on the 0.07 mm threshold. The potential for sexual

reproduction is reflected by the observations of a 1:1 sex ratio found at all recreation

sites and throughout the field season. I also found that 10% of the males surveyed

showed ruptured sperm follicles, which indicated that there was adequate nutrition for

spermatogenesis and that sperm was released into the water column at the time these

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males were collected in mid-May to early June 2011. In contrast, all eggs were not

sexually mature based on Littleford’s (1939) egg diameter maturity index of greater than

0.07 mm and because no brooded planulae were observed in the female C.

quinquecirrha sampled. Based on these results, I did not have sufficient evidence to

prove that sexual reproduction had occurred. Mature gonad colors of female C.

quinquecirrha in Chesapeake Bay were described to be grayish-yellow brown and

males to be bright pink (Littleford 1939). Although my color comparison of female and

male NRE C. quinquecirrha did not coincide with Littleford’s observations, it could be

that mature gonad colors were not observed because the majority of the jellyfish were

indeed sexually immature. However, as documented in other jellyfish species (Schiariti

et al. 2012; Toyokawa et al. 2010), it is also possible that gonad color is unrelated to

mature gonads. Based on my results in chapter 3, I cannot attribute sexual reproduction

of C. quinquecirrha as an ecological driver of jellyfish-human interactions in the NRE.

However, it is clear that more research is required to fully support this conclusion.

Planulae settlement studies within the oyster sanctuaries and on docks throughout

APES would aid in determining to what extent sexual reproduction contributes to the

annual occurrences. These observations coupled with histology, in vitro fertilization

experiments, and genetic surveys would give us a better understanding of the

reproduction cycle of C. quinquecirrha in APES, which would help with managing this

species.

Cultural consensus analysis found that perspectives of jellyfish ecology varied

among the four cultural groups, but societal perspectives were similar. The cultural

perspectives of jellyfish researchers were closely aligned with what is biologically known

176

about jellyfish, but perspectives on jellyfish ecology among the other groups were less

clear. Specifically, there seems to confusion on the role jellyfish play in food webs, as

jellyfish were placed in lower trophic levels by fishers, recreationists and 39% of the

coastal researchers. Improper placement of jellyfish in food webs could be due to an

“ecological misconception” (Cherrett 1989), which stems from the participant’s prior

knowledge and experience with jellyfish. This supports the argument that the lack of

data and education about jellyfish in ecosystems has reverberated to the research and

public interfaces. Therefore, one of the social drivers that influence jellyfish-human

interactions is confusion in jellyfish ecological literacy.

Regarding the influence of jellyfish on society, all groups share similar

perspectives that jellyfish may be problematic to coastal communities, but that socio-

economics may not be affected due to varying amounts of tolerance toward seeing

jellyfish in water and stings. For example, seeing fewer than five jellyfish in water was

perceived to be tolerable because all groups agreed that this amount would not stop

water activities. There was a notable difference in perspectives when the groups were

asked if water activities would cease if more than five jellyfish were present and this

could be a reflection of the jellyfish species encountered (i.e. the cannonball jellyfish S.

meleagris versus the sea nettle C. quinquecirrha) and the type of water activity the

people are engaged in (i.e. fishing, surfing, researching). Moreover, despite the

negative repercussions of jellyfish stings, all groups agreed that people would take a

vacation to areas where jellyfish are sighted regularly even if they are stung by jellyfish.

Based on these data, I conclude that another social driver that influences jellyfish-

177

human interactions is the acceptance of certain amounts of jellyfish in the water

regardless of the severity of jellyfish stings.

Since the cultural perspectives of jellyfish in food webs were less clear among

fishers, recreationists and coastal researchers, to improve jellyfish ecological literacy,

outreach education that includes the role that jellyfish play in food webs should be

conducted and coastal researchers should be encouraged to include jellyfish in their

studies. In the NRE or other areas within APES that are prone to C. quinquecirrha

occurrences, the use of barrier nets to protect swimming areas could prevent harmful

encounters as demonstrated in Chesapeake Bay (Schultz and Cargo 1969) and

Barnegat Bay, New Jersey (Vasslides and Sassano 2012). As well as preventing

contact with C. quinquecirrha, the barrier nets could reduce the number of C.

quinquecirrha to the fewer than five tolerances, which could prevent decreases in

estuarine recreation and subsequently tourism revenue would not affected.

Residents reported that they enjoyed seeing and swimming with the cannonball

jellyfish S. meleagris off the coasts of Oak Island and the Outer Banks (Figure 1). This

type of interaction may be similar to swimming with Mastigias sp. in “Jellyfish Lake”,

Palau. Moreover, jellyfish are aesthetically appealing in nature and also in aquariums

worldwide. Fishers will use the cannonball jellyfish S. meleagris as bait for various types

of fish, including spade fish and residents that frequently use the beaches of Oak Island

(Figure 1) told me that they would be sad or “miss” seeing S. meleagris in the water if

these jellyfish were to suddenly disappear. These interactions could render the

classification of a positive jellyfish-human interaction. Furthermore, since S. meleagris is

a commercial species that is heavily fished in Georgia, the potential utilization of S.

178

meleagris for North Carolina fishers could be beneficial. To assure the viability of a S.

meleagris fishery in North Carolina, proper assessments of abundance and research on

how the cannonball jellyfish affects the overall coastal ecosystem must be determine a

priori.

Residents also have encounters with the “non-stinging” ctenophore comb jellyfish

M. leidyi. Encounters with this species do not harm humans but residents observe M.

leidyi quite often. North Carolina residents will refer to M. leidyi as the “moon jellyfish.” I

was surprised to hear that the common name of “moon jellyfish” is used to describe M.

leidyi because this common name is usually associated with the cosmopolitan Aurelia

sp. jellyfish. Future jellyfish research in North Carolina should be aware of the use of

this common name, especially if discussing jellyfish with local residents. Lifeguards and

beach-goers have observed the presence of the sea nettle on the Outer Banks beaches

(Figure 1) each summer. Since the sea nettle is an estuarine jellyfish species (Cargo

and Schultz 1966) it is possible that the sea nettles observed on the Outer Banks

beaches were transported by an alongshore current that passes by the mouth of

Chesapeake Bay. Although the gelatinous bodies of jellyfish poise problems for tagging

experiments (Purcell 2009), set-nets and gonad maturity assessments were used to

track the growth and movement of the giant jellyfish N. nomurai in the East China Sea

and Sea of Japan (Toyokawa et al. 2010). Remote sensing or aerial photography could

be used to observe if the sea nettles found on the Outer Banks originate from

Chesapeake Bay. If these sea nettle occurrences are found to originate from

Chesapeake Bay, the NOAA sea nettle “nowcasting” model (NOAA 2013) could be

used by Outer Banks residents and tourists to prevent unwanted sea nettle encounters.

179

Evidence that jellyfish populations have increased in varying amounts of severity

over the past century and the anticipated development of coasts worldwide suggests

that jellyfish-human interactions are likely to continue. To manage these interactions, I

have learned that multifaceted research practices provided substantial insight to what

drives jellyfish-human interactions in North Carolina. I believe this approach would

benefit other areas prone to jellyfish occurrences in lieu of the singularity of traditional

jellyfish biology and/or ecology studies. To date, ecosystem-based management has

encouraged researchers to expand their research scope from single species to studying

the complexity of ecosystem dynamics (Rosenberg and McLeod 2005). Although this

was a step in the right direction, incorporating human dimensions into ecosystem

research has been difficult because of the extended amount of time between practice

and policy implementation (Pikitch et al. 2004; Smith et al. 2007). It is hoped that current

research endeavors using transdisciplinary science, where hypotheses are created with

specific goals that benefit the sustainability of society (Lang et al. 2012), will curtail

disparities between research, policy, and the general public. Under this auspice,

scientists are encouraged to seek collaborations with colleagues from different

academic backgrounds to investigate research problems that are important to the

general public. By studying what drives the multifaceted interactions that surround

jellyfish and humans, future studies will continue to benefit the sustainability, utilization,

and management of coastal resources influenced by jellyfish-human interactions.

180

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quinquecirrha). Hydrobiologia 451:89-95.




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Lang, D., A. Wiek, M. Bergmann, M. Stauffacher, P. Martens, P. Moll, M. Swilling & C.

Thomas, 2012. Transdisciplinary research in sustainability science: practice,

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Mallin, M. A., 1991. Zooplankton abundance and community structure in a mesohaline

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Administration, Chesapeake Bay Office.

http://chesapeakebay.noaa.gov/forecasting-sea-nettles Accessed 20 March 2013

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Paerl, H. W., J. L. Pinckney, J. M. Fear & B. L. Peierls, 1998. Ecosystem responses to

internal and watershed organic matter loading: consequences for hypoxia in the

eutrophying Neuse river estuary, North Carolina, USA. Mar Ecol Prog Ser

166:17-25.

Pikitch, E. K., C. Santora, E. A. Babcock, A. Bakun, R. Bonfil, D. O. Conover, P. Dayton,

P. Doukakis, D. Fluharty, B. Heneman, E. D. Houde, J. Link, P. A. Livingston, M.

Mangel, M. K. McAllister, J. Pope & K. J. Sainsbury, 2004. Ecosystem-based

fishery management. Science 305:346-347.



182



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Purcell, J. E., D. L. Breitburg, M. B. Decker, W. M. Graham, M. J. Youngbluth & K. A.

Raskoff, 2001. Pelagic cnidarians and ctenophores in low dissolved oxygen

environments: a review. Coastal and Estuarine Studies:77-100.

Rosenberg, A. A. & K. L. McLeod, 2005. Implementing ecosystem-based approaches to

management for the conservation of ecosystem services. Mar Ecol Prog Ser

300:270-274.








Shoji, J., T. Kudoh, H. Takatsuhi, O. Kawaguchi & A. Kasai, 2010. Distribution of moon

jellyfish Aurelia aurita in relation to summer hypoxia in Hiroshima Bay, Seto

Inland Sea. Estuarine, Coastal and Shelf Science 86:485-490.

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Stanley, D. & S. Nixon, 1992. Stratification and bottom-water hypoxia in the Pamlico

River estuary. Estuaries 15:270-281.





Thuesen, E. V., L. D. Rutherford, P. L. Brommer, K. Garrison, M. Gutowska & T.

Towanda, 2005. Intragel oxygen promotes hypoxia tolerance of scyphomedusae.

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Vasslides, J. & N. Sassano, 2012. Early results from the Barnegat Bay Sea Nettle

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184

LIST OF FIGURES

Figure 1. Map of eastern North Carolina, including the Albermale Pamlico Estuarine

System; Albermarle Sound, Pamlico Sound, Pamlico River Estuary (PRE), and the

Neuse River Estuary (NRE).

185

Figure 1.

186

Appendix A: IRB letter/approval for Jellyfish Survey

EAST CAROLINA UNIVERSITY University & Medical Center Institutional Review Board Office 4N-70 Brody Medical Sciences Building· Mail Stop 682 600 Moye Boulevard · Greenville, NC 27834 Office 252-744-2914 · Fax 252-744-2284 · www.ecu.edu/irb

Notification of Exempt Certification

From: Biomedical IRB To: Mahealani Kaneshiro-Pineiro CC: Hans Vogelsong Mahealani Kaneshiro-Pineiro Date: 5/10/2012 Re: UMCIRB 12-000609 Jellyfish survey I am pleased to inform you that your research submission has been certified as exempt on 5/10/2012. This study is eligible for Exempt Certification under category #2. It is your responsibility to ensure that this research is conducted in the manner reported in your application and/or protocol, as well as being consistent with the ethical principles of the Belmont Report and your profession. This research study does not require any additional interaction with the UMCIRB unless there are proposed changes to this study. Any change, prior to implementing that change, must be submitted to the UMCIRB for review and approval. The UMCIRB will determine if the change impacts the eligibility of the research for exempt status. If more substantive review is required, you will be notified within five business days. The UMCIRB office will hold your exemption application for a period of five years from the date of this letter. If you wish to continue this protocol beyond this period, you will need to submit an Exemption Certification request at least 30 days before the end of the five year period. The Chairperson (or designee) does not have a potential for conflict of interest on this study. IRB00000705 East Carolina U IRB #1 (Biomedical) IORG0000418 IRB00003781 East Carolina U IRB #2 (Behavioral/SS) IORG0000418 IRB00004973 East Carolina U IRB #4 (Behavioral/SS Summer) IORG0000418

187

Appendix B: Jellyfish survey consent form

Study ID:UMCIRB 12-000609 Date Approved: 5/10/2012 Does Not Expire.

Jellyfish Survey Participant Agreement

You are being invited to participate in a research study titled “Jellyfish-human

interactions in North Carolina” being conducted by Mahealani Kaneshiro-Pineiro, a PhD

candidate at East Carolina University in the Institute for Coastal Science and Policy,

Coastal Resources Management PhD Program. The goal is to survey 150 individuals

in/at beach, coastal and estuarine recreation areas and research institutions in North

Carolina as well as jellyfish researchers located primarily throughout the United States.

The survey will take approximately less than 30 minutes to complete. It is hoped that

this information will assist us to better understand what people know about jellyfish and

how jellyfish may influence a person’s choice to engage in water activities. The survey

is anonymous, so please do not write your name. Your participation in the research is

voluntary. You may choose not to answer any or all questions, and you may stop at

any time. There is no penalty for not taking part in this research study. Please call

Mahealani Kaneshiro-Pineiro at (252) 328-9375 for any research related questions or

the Office for Human Research Integrity (OHRI) at 252-744-2914 for questions about

your rights as a research participant.

188

Appendix C: Jellyfish survey

Contact: Mahealani Kaneshiro-Pineiro Phone: 252-328-9375

[email protected]

Instructions:

Please answer all questions by yourself and to the best of your ability Do not look up any answers to the survey questions If you do not know the answer to a question, guess

If you have any questions while taking this survey, please contact Mahealani for assistance

189

1. When was the last time you went to the coast/water?

____________________________________

2. Did you see jellyfish?

Yes No

3. Have you ever been stung by a jellyfish?

Yes No

4. What months do you usually do your water activities?

____________________________________

5. On average, how many hours per week do you spend doing your water activities?

____________ hours per week

6. Where do you go for your water activities?

____________________________________

7. Are jellyfish usually in the water when you do your water activities?

Yes No

If you answered YES to the previous question, please answer these next questions:

Are jellyfish new to your coastal waters, beaches and/or shorelines?

Yes No

Over the past 5 years, have you seen more jellyfish in the water and/or on shorelines?

Yes No

Over the past 10 years, have you seen more jellyfish in the water and/or on shorelines?

Yes No

Look at each picture and please write down the name of each in the space below.

_____________

_____________

_____________

_____________

_____________

Please write any other names you know here: _________________________________________________________

Please tell us about your experiences with water activities and if you have seen jellyfish while doing your water activities.

190

For these questions, please tell us if people agree or disagree with the following statements about jellyfish.

01. Jellyfish are animals Agree Disagree

02. Jellyfish are plants Agree Disagree

03. Jellyfish live longer than a year Agree Disagree

04. Jellyfish have eyes Agree Disagree

05. Jellyfish have a brain Agree Disagree

06. Jellyfish have a heart Agree Disagree

07. Jellyfish use fins to swim Agree Disagree

08. Jellyfish use tentacles to swim Agree Disagree

09. Jellyfish can swim against moving water Agree Disagree

10. Jellyfish thrive in murky water Agree Disagree

11. Jellyfish do not need a lot of air-in-water to survive Agree Disagree

12. Jellyfish need to come up to the surface to breathe Agree Disagree

13. Jellyfish come up to the surface to eat Agree Disagree

14. Jellyfish eat fish Agree Disagree

15. Jellyfish eat shrimp Agree Disagree

16. Jellyfish eat plants Agree Disagree

17. Jellyfish can eat more than their body weight Agree Disagree

18. Jellyfish do not need to eat to survive Agree Disagree

19. People eat jellyfish Agree Disagree

20. Turtles eat jellyfish Agree Disagree

21. Dolphins eat jellyfish Agree Disagree

22. Fish eat jellyfish Agree Disagree

191

For this question, think about jellyfish and their importance in nature. Rank the following #1 to #9,

with #1 being the most important in nature and #9 being the least important in nature.

Which food web picture or pictures best describes jellyfish in nature? Please circle your answer(s).

______ ______ ______ ______ ______ ______ ______ ______ ______

192

For this question, think about how people perceive jellyfish; specifically, think about the fear of jellyfish and how that would compare to other living creatures.

Rank the following #1 to #9, with #1 being the most feared and #9 being the least feared.

______ ______ ______ ______ ______ ______ ______ ______ ______

For these questions, please tell us if people agree or disagree with the following statements about jellyfish stings.

193

How many jellyfish seen in the water will make people stop their water activities? Please circle your answer.




194

For this question, think about the aquatic ecosystem. How do you think people should spend science money?

Rank the following #1 to #9, with #1 being the most money and #9 being the least money.

______ ______ ______ ______ ______ ______ ______ ______ ______

For these questions, think about jellyfish and society. Please tell us if people agree or disagree with the following statements.

195

Please answer the following questions about yourself. The information you provide will remain strictly anonymous and your name will never be associated with your answers.

1. Are you (circle one)

(a) Female

(b) Male

2. What is your age?

__________ Years

3. Which state and country are you a resident of?

____________________, _______________

I4. Which category best describes the highest level of education you have completed? (circle one)

(a) Some high school; grade completed _____

(b) High school/equivalency

(c) Associate’s (2 year degree)

(d) Bachelor’s (4 year degree)

(e) Graduate

5. Which group best describes you? (circle one)

(a) Fisher

(b) Coastal recreationist (swim, ski, surf, etc.)

_________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________

Survey ID#_____________

In the space below, please share anything else you know about jellyfish. For example, when do you see jellyfish? Or use the space below to write any other comments about this survey. Thank you!

196

Please answer the following questions about yourself. The information you provide will remain strictly anonymous and your name will never be associated with your answers.

1. Are you (circle one)

(a) Female

(b) Male

2. What is your age?

__________ Years

3. Which state and country are you a resident of?

____________________, _______________

I4. Which category best describes the highest level of education you have completed? (circle one)

(a) Some high school; grade completed _____

(b) High school/equivalency

(c) Associate’s (2 year degree)

(d) Bachelor’s (4 year degree)

(e) Graduate

_____________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________________

Survey ID#_____________

In the space below, please share anything else you know about jellyfish. For example, when do you see jellyfish? Or use the space below to write any other comments about this survey. Thank you!

Documents

by Mahealani Y. Kaneshiro-Pineiro Dissertation Advisor ...thescholarship.ecu.edu/bitstream/handle/10342/1745/...Dissertation Advisor: David G. Kimmel, PhD Major Department: Coastal